THE TERMITE FAUNA. (lSOPTERA) IN THE VICINITY OF

i)
from Tcrtiary Ambcr of
THE TERMITE FAUNA. (lSOPTERA) IN THE VICINITY
OF CHAMELA, STXrE OF JALISCO, MEXICO
e Psocoptcta. Jlcmoir 14.
ity Statistics aml tite Log
Distriilution of A.rbOical
:kroachcs. Occologia (/Jcr-
DA\'IIJ
l'\ICKLE
.\.
\[A.RCARLT
*
S. COLI.INS
**
* Sy,tcmatic Entomology
Lahoratory
USD.\ t:/o U.S. National :\fuseum
of l'\atmal Hi'itory. NHB-16H
\Vashington. D. C. ~0560
* * Dcpartmclll of Entornology
U.S. ~ ational Musen m of
1\atural History. NHB-168
Washington. D. C. 20560
U.S.A.
Folia Fnlo111olágica Mexicana .\n. í7: 85-122
RclCi\ ed for publicalion: Septcmher llJ. 1'ISS .
. \cccpted for puiJlication: Dcccmbcr ~0. El~S.
8:í
(l9HH)
86
FoLIA
E:--:rmiOL. :\IF:X. 77 (1988)
:\ICKI
The termite fauna of thc dry scrub forcst region of Jalisco State, l\fcxiw. is
rcprescntcd at thc Chameb Estación, l. :'\AM, and nearby arcas by more than 27
'pecics. Ten specics of dry-,rood tcrmitcs iu the genera 1\'cutermcs, Jncisitcrmcs. Jfarginitcrmes, Calcaritennes, and Cry¡Jtotermrs (Kalotermitidae) are common in thc
hanlwoods ami timber. Although only two subterraucan specics, Coptotermes cr11.11tts
~nyder 1922 and Heterutermes aureus cont~exinotatus Snydcr 192-4. rcprescnt the familv
Rhinotermitidac, thcy constitute thc 1110 potentially major pcst specics of thc :nea.
Fiftecn spccies (comprising thc genera Ho¡Jlotermes, AnofJ/otcrmes, fo.Iicrocerokones,
Amitermes, Gnathamitermes. Termn, Sasutitermes, and Tenuirm!ritennes) of thc most
advanced familv. thc Termitidac, are a major componcnt of thc inscct fauna of thc
rcgion, in tcrr~s of numlwrs of in di 1 iduals and numbcrs of termite specics.
KEY '\'ORDS: Isoptera, Ta~onomy, Drv scrnb forest, Jalisco,
.vlcxico.
La far111a de termitas dd bosque caducifolio de Jalisco, '\léxico, está bien L<"l" ,._
scutada en la Estación Chamela, UNAl\f. v en las áreas ccrctllas, dPJHle se encuentrau
m;ís de 27 especies. Die? especies rclacic;nadas con las maderas secas, in el nidas en
los géneros Neotermes, Inci;itcrmes, J!arginitcrmes, Ca/caritcrnu:s y Cryfilotennn (Kalotcrmiticlac) son comunes en los árboles con madera dura. Sólo dos especies subten:meas,
Coj>lotermes crassus SnHh r I~J22 y Heterolcrnzcs aureus convcxinolatus Snydcr 1~J24
1Rhinotermitidae) tienen importancia como plagas pott:ncialcs. Las 15 especies de
Tcrmitidac incluidas en Jos gtneros Hoplotcrmes, Anuplotermes, 1\Iicroccru 1crmcs,
.·lmitnmes, Grzatlzamitermes, Termes, l\'asutitermes y Termirostrikrmn, re¡ll("eutan
un componente importante de la entomofauna de la región, tanto por su número ele
indi\"iduos como por la cantidad de especies.
!'.\LABRAS CL\VE: Isoptera. Taxonomía. Bosque Tropical Dclucifolio. Jalisn• . .\h'xico.
l!'\TRODCCT!O'\'
Thc tcrmitc fauna of Chamela and a<ljacent :u·cas of the St:ltc of
Jalisco, ;,rc:\:Íco, i-; divcrsc. combining a fcw Ncarctic: cxtcn:;ion'i 11·i¡)¡ thc
more mnncrous :'\t:otropical clemcnts. "\t lc;,,t twenty-se,-cn spccies are
rcp1 co;c¡¡Lcd lJcJo¡¡g;ng to thrcc tcrmitc familic-, - J.;.alotcnnitidae, Rhinotcrmitidac, ami Termitidac. 1nclmling woocl-dwclliug, ~ubterrancan, and
carton-ncst building
eco loo ical ancl beh;
"'
')rimitiYe
termite~.
l
lonies in the deacl w
Some are imponant
specic-; of this biol
rare in thi-; rcgion
these tcrmites, as "
important cnYironn
trient rccycling, by
nents aYailablc for
\Iunbers of the f
11·oocl-ch\·elling ~pec
nutrient-recycling a
tentially scriom pe
t he arca, H ctcrutcr
tcrllics crassus Sny<
pc'>t species for th
Thc most ~pecializ
,pecic~ in eight gc
mouncls or conspic
are also tbe most
thc tropics.
\\'e fin el that arr
intere;,t in identifi1
ldentification of s
tification keys inc
lJ;¡ ,e, of onc or a
not ;n ailable. Fun
this p<tpcr) had be
was latcr describec
t ity of a ,pccies ;
umlrum. Tc1mite
phological feature'
rcsohing termite
char;1cter for "pec
;tgc of the colon¡
mctn, also yary
n:',ult~ from thc
m lll: Jh,, with oldc
a¡ ti,mcs has rcsul
~)
:'\tcKLr: ET CoLLI:\S.
Jalisco Statc, l\fexicu, is
arcas by more than 27
~ermcs, Incisitermc;. Jfare) are common in thc
des, Coptotermes crrrsms
924. rcpresent th.:: famih·
pcst specics of the ;:¡ 1e;;.
ilcrmes, 1\licrocerokones.
rostritennes) of thc most
the insect fauna of the
ter mi te species.
~.
;\Icxico.
;féxico, está bien 1cp; eas, donde se encuentr;m
eras secas, incl nielas en
·s y Cryptotermcs (Kalolos especies suhtcn;inca 'rexinotatus Snydcr ¡~)'>4
es. Las 15 e¿pccics
~ernzes, J\.Iicroccru.fcnncs,
•ostritermes, repre;eutan
anta por su número de
de
:aclucifolio. Jali,n·. '.f<'-
~as
of thc Statc- of
extensiom \1-itlt thc
1ty-sen:n spccies are
lotermitidae, Rb in u' subterranean, ancl
TER:'IllTFS oF CHA::\IEL\
87
canon-11est building -,pecie-;, they illmtrate sorne of the great diversity in
ecological ancl behavioral characteristics found in termites. The more
primitive termites, belonging to the family Kalotermitidae live in colonie'> in the deacl wood of living or dead trees, fence posts, and buildings.
Sorne are imponant pe'>~' of structural timber. On the other hand, severa!
specic:-; of this biolugitally interesting group are becoming increasingly
rare in thi-, rcgion as wood usccl for fuel is consumecl by man. l\fany of
these tcrmites. a' well as members of the othcr two families, play an
important envirunmcntal role ·within the community as agents of nutrient rccyding, by breaking clown clead "·ood and making its components aYailahlc for liúng plants and other animal species.
\lt::mber<; of the famih Rhinotenniticl:tc include both subtcrrancan ancl
wood-d\\·elling '>pecie-,; unfortunately, although they are also important
nutricnt-recycling agenh in natural environments, most species are potentially seriou-, pcsh of structural timher. The two species known from
the areC\, H ctcrutcrmes aurcus cunvcx inotatus (Snyder) 1924 and Captoter/l/ es crassus Snyder EJ22, are common aml constitute the two major
pe<;t species for this rcgion, frequently attacking man-made structures.
The most .'pecialized family, the Termitidae, is representecl by fifteen
>pcrie'i in eight genera. Most of the species \\'ithin this family build
mouncls or conspicuous carton nests on trees, stumps, and post'i. They
are also the most numerous of thc tcrmites both here and else>vherc in
the tropics.
\\'e find that among other rescarchers tbcre is a growing need for and
intere,t in identification keys to clifferentiate species, based on all castes.
Iclentification of specics of termites can be clifficult. In the past, identification key~ incluclecl some spccics thC\t had been describecl on the
b;,,e, of one or a fe\r specimens, aml detailecl illustrations were often
not aY;tilahlc. Furthermore, many spccies (including severa! reviewed in
th is paper) had be en dcocribecl from only one caste, while thc other e aste
was la ter described as yct anot her species, casting· cloubt as to the iclentity of a species aml thereby re:-.ulting in a confusing taxonomic conumlrum. Tcnnite specics instead shou]J be charactcrize<l basecl on morphologic:tl features of both solclier and imago castcs. 1\nothcr problcm in
rcsohing termitc idemifications has bccn the Lictor of using size as a
ch;m:cter for '>¡wcies recognition. Si?e, c<;pccially in soldiers . yarics with
;tge of thc colcny, and frcqucntly ratios hetwccn morphological paramctLT:-. aho Yary ~~·ith changcs in oize. \'ariahility in soldier size oftcn
rc.·,ults fmm thc fact that 'olcliers can originate from different instar
nym;Jb, \\'ith oldcr nymphs producing larger soldicrs. This phenomcnon
at times has resultcd in misiclentification of '~pecics or even in thc de-
SR
FoiL\ E:\TCJ:\IOL.
:\In. 77 (19HK 1
scription of t\\'O separatc -,pecies. ln futurc -.,tudie' thc,e problcm., may be
rcsolwcl if care is taken to obtain larger ~amples from indiYidual colonies
and if colonies are maintainccl in thc laboratorv or monitored in the
field until winged adults are gencratcd.
that both rcproducti,·c all(l
soldicr ca:,tes can be associatcd for clcKriptions.
In this paper we combine the information \\'C acquircd on two field
trips to we-,tern :\Iexico (1%1-i by :\fSC ami l~li' 1 h; D.-\~ ami .\TSC)
with the information publishcd h; Light (1933) ami Sn;clcr (1922, 192·1.
1946). \Vherc possible, \\'C prm iclc bio1ogical infonuation. ;me! wc prescnt identification keys ami figure-, to Ltcilitatc rccognition of all spccies known to occur in thc region. Unfortunatcly. sincc thc aLttcs of
fourteen species are unknmrn, it ha-; not heen po"ihle Lo construct an
adeuuate imago identification key for all o[ the bun;¡ of the regiotJ. Then.:forc we present a simplifiecl imago kcy to thc le\·el of gcnus. \\'ith the
expectation that continucd research on termitc' Hil lill in thc gaps of
our current knowledgc of thc-,c life fornh.
~!CK
-.o
PROCLlll 'Rb
Collccting tennitcs. Currcmh· there are onh ;j lc\1· actÍ\'C termitc
spccialist'i im·olvcd in termitc (üllccting, ancl mtr-<.:um ancl uni,·ersit;
holdings are depaupcrate in severa! groups. Thc fo!lowiug information
i~ provicled for futurc termite collcctors in the ;trea. Carton-ncst building
termites wcre ohtained by cutting into cithcr nmwa}" or the nest proper.
prefcrahly \\-ith a hea\) blacled machete or hand axe, Gtlching thc
fragmenh in a tray, :me! removing thc termitc-; inclividualh·. Spccimcm
to he kcpt :tlive werc handled with a papcr spatula (Fi::\·· ]) rather than
forceps so a-, not to damage cutide. To en,m·e that ,¡wcimcns to be
kcpt alivc woulcl not be stres-,ccl or injured. thc hbdc \LlS ~lid gcntly
tmdcr the fect of the -;pccimcn, \\'hich mu;tl!; dung to ir until tapped
into a container. If the ;mim:tl:, offcred to hite the hlade. a frcsh spatula
\\':ts used.
Larger ';11nplc, were obL1 i ned rapic!Jy by u-,i ng p;t i red -,ic\ e,, one \\'ith
rclat.ively brgc opcuing'-> (:\'o. 10 Standard, ~.O mm) ovcr une \l·ith '>mallcr
(:\'o. 1~ SLilltl:ird, l.OmmJ opcning-.. \\'!ten tllc ne-;¡ lr;!gmulh \\·ere phccd
in thc hrger-mcsh ,¡c,·c. thc lcrmitc-, migLttcd dmvm1·;ud. \1·herc thcY
wcrc c;¡ught in thc smallcr-mc-;hed couLtincr. l'crmitopb!lc-; \HTe collectccl
by cuttin:~ tlw inner portion, of tbc lll''it into clnmk,, :illtm·ing thcm to
sit for a ,JlOrt time. then sL11nming the nrt -;urLice of thc nc,t fragmcnt
ag:1imt tlw floor of a \ellm\·-colored pLhtic Lray (of thc t\pc freqncnth·
r-------!--1
-
-
- - - -r--;
lig. l. .\ paper spatul
-,p:nula is constructed by
thc rcmainder along the
,o]d for th<: ;" a dio
of thc termitc "gtH
mar; rcproductivcs
\l'erc ahray-; collccter
Su btcrr;¡uc:m tenr
ti:ilh-huricd decayin
,uft or rotten, a hcar
l\ pe. G:tllcrie-. in in
di'J:Jdgcd by gcntly
,jC\C\ 'I'CI'C mcfuJ f,
{;r;¡"-:md humus-fccr
r ow d1mg. do11·n wo
quickh to greater de
i 11;_: a t ro m:' l or small
mcdiatdy along witl
g:d lcrie-,. Tcrmitcs ;n
trca \1-:~'> '>Carchecl fo
fo1 'cparating the te
1)¡ Y-m)()d Lcrmitcs
,;,-,.-_ ;m ::\.e or hatch
,,
')
l'\ICKLE
FT
CoLu:-.;s.
TER:\IITEs oF
CHA:\HLA.
lbe problem~ may be
m individual colonies
or monitored in the
~th reproclucti,·e allC!
cquired on two fielcl
by DA:\' ancl .\fSC)
I Sn;der (1922, l92i.
mation. ancl we precognition of al! spesince thc ;tia tes of
;ihle to construct an
of tl1e region. There1 of genus, with thc
' fill in thc gaps of
fe11· acti\'e tennite
eum ancl uni,·ersit\
•llowing informatio11
Carton-nest huilding
; or the nest proper.
axe, catdting thc
iviclually. Specimcn~
(Fig. l) rather than
at specimens to he
acle was slid gently
: to it until tapped
acle. a fresh spatub
red sie,·e,, one with
¡er one 11·ith .,mallcr
tgmcnt., wcre pbccd
·nward. 1dwrc theY
>hile-; "·ere collectcd
;, :lllmring them to
E the ne.,t fragment
the type frequentlv
- - - - - -1-----------------------t
- - - - - -1----------------------f
1· i¡;. l . .-\ paper spatula for handling liH~ tennilcs withoul damaging cuticle. Thc
·,pattda is comtructed by cutling a :l x '>" imlex can! along tia· dotted Iines ami folding
lile rcmaindn a long thc sol id iines LO !onn a handlc.
,oJd for use a-, ;¡ dishpan). This light background lacilitatccl recoglutton
of the tcnnite "guests", typically found near thc royal chamber. Primar; rcproducti\cs also wcrc found in this m;tllner. If present, aLtte-,
1rcre alway, collened first to as'iurc association of alf caste-, for a sample.
Subtcrr;tucm termitcs frcquently were founcl in fallen, buried, or parti;tlh-huricd decaying wood. In working with woocl that might Le fairly
.,oft or rotten. a hea,·y machete w:ts more uscful than a thin-bladed sharpcr
t\ pe. Galleric, in infcsted wood werc opened carefnlly :md the terrnitc'
di.,Jaclgcd by gcntly tapping thc exposccl wood o1·cr tray-; or .,icve"i. Thc
,icye-; \\·ere mcful for separating the tennites from the nest fragmenh.
Cr;"s-aud humus-fccding .,ubterrancan tennites werc locatcd under stonc'.
cm1· dung. down ,\·ood, debri.,, cte. Once exposed. these tcrmitcs rctreat
quickly to greater depths, so it was usu:dly necessary to collect quickly. Esin;.; :t llO\ITI or .,mal! shon·l, samplcs of cxposcd termites "·ere g:tthered immcdiatcly along with a cut o[ soil hcfore thcy could retrcat to decpct
g:tilcrics. Tcrmite., ancl soil werc thcn placed in a tr:l\ "·hile the :tdjaccnt
:trc;¡ \l':t' scarchcd for additioual 'ipccimen-; .. \g:tin, tlw 'iÍC\'CS "·ere u-;eful
íot 'cparating thc tetmitcs from ucst material.
Dn-wood l<Tmitc.;; werc found in '>Ouncl woocl, requiring t!Je use of a
,;¡w, ;~n ::xc or l!atchct, ancl a he:ny-hladed machete .. \ qout :o.crewdrin'r
90
FollA E;-.;T0\10L. :\II:x. 77 (ElRR)
\\·as mcful in opening inhabited galleries, from which the termites could
Le gently tapped into smooth-~urfaced basins or trays. Since these termites
usually dung to nest fragmcnts rather than attempting to trawl down"·arcl, ~icn:s were not meful for collecting most dry-wood termites. ln'>tead, incliYiduals hacl to be transferred to Yials with specimen forceps,
or with spatulas to collecting chambers if the specimens were to be
kcpt aliw. H al ates were present, they were harvestccl q uickly, sin ce disturbanccs to the nest usually inYoked flight bchaYior. If reproductiYe
nYrnphs of aclYanced instars were prcsent but aL1tes hacl not yet cleYeloped, it ,,-as sometimes possible to rear the nymphs to adulthoocl to obtain
winged forms. Samples of workers ancl uninjurcd termite reproductive
nymph-; we::re provided with food, moisture, ami protection from cxcessin:
heat, clirect 'unlight, ancl predators. Vragmcnts of ncst material aho wcrc
incluclnl.
Prcscrvation of samples. Specimcns were preserved in 85'; ;. cthanol, \l·hich
was replacccl after 24 hr, especially if a large samplc werc ohLtincd_
Data on the Jabel included locality, date, name of collector, type of hahitat; if alates wcre taken during a flight, the time and weathcr conclitions
preceding ~md dming the flight were also notcd.
.'\f aterials and mcthods. This study was ha sed on specimcm in the N ational
~1uscum of ;'\atural History, vVashington, D. C. [N \!NHJ, and thc
American :\1useum of Natural History, New York. :NY [A\INH]. Some
of the spccime::ns collectecl by us "·ere donatcd to the Estación de Biología,
Chamcla [CHAMJ and to the .\luseum of Comparative Zoology, Hanarcl
UniYcrsity, Cambridge, ::\ rA [;\ICZ]. Spccimcm U'iCd in thc scanning elecaon micrographs (SF:\Is) were those collcctcd by Nickle and Collins in
198~ in e u:ttemala and l\lcxico as part of their ficld studies for a book
in prcparation on the termites ol North America. Spccimens to be photomicrographcd '1-cre fir-;t transfcnccl to vials of absolutc ethanol ancl
maintainccl at this conccntration for 21 hr prior to drying. They ,\.el-e
thcn clried in a S.\ .\!D RI critica! point drier using· teclmiqucs describccl
hy Gonlh ;md Hall (197~J). Driccl spccimem \\-ere fastencd \\'ith Elmer's R
gluc cither to a paper point or dircctly into ;¡ metal stub for used in thc
'cann~ng clenron microscope. They wcre thcn gold-coatcd, cxamincd, and
photomicrohraphcd at 1:) 1~ yo]ts, ming a Cambridge Scanning Elcctron
\!inu wpc at an a]-'!HOpi iatc magníficat ion to show spccie, diffcrellccs.
Spccimcn~ til;~L conld nct be stucl~ccl \\·ith an SE\!, eithcr bccamc thcy wcrc
in slw: t 'u:Jph or ya]uablc a' unitluc :,:>Ccimcm. werc rcndercd while im-
.\;ICK
II!Cl'SCd
in
8:J<·~ Ct!J
e amera 1u cicla.
::\early all figuree
~pecies, reponed in
from spccimens ot
imagoc-, of Xeoten
, Haldeman) 1853, a
eomiclerecl that for r
to illustrate intersp
'ic11·, . .\Iost of the
im:1gc' for the firs1
'pecic, recognition
in thi.> rcgion.
Thc llames of tl
listccl by R. L. Ara
·.pcrimem. especia]]
SPJCU:s OF
K.~LOTlR\llTIDAE
Scotames joute/i (E
lncisitcnnes
ni{!:ritus (Snydcr)
111a1 ginijiwnis
(L
jJ/a/\1 cpiwlus (Li
tlllCf.IUIIÍ ¡Light)
,\Iarginitn mes hubba
Calraritcrmcs
jHm>inolll\ (Light
!rmgirollis (Banks:
(;¡ Yjllolcrmcs
(atulus
lm·vis
ILight) l!
1
1 Walker)
Rtii~OlTK.\ílT(]) \E
llctcmtn mes aurc
Coj,ín!crJJ/r'(\ crasq
.·lnu/'loicrnu:s fu mas u
l)
:\IcKLE ET CoLLI:\s.
ich the tennites could
rs. Since these termites
pting to travel downlry-wood termites. Inlith specimen foneps,
pecimens wcrc to be
tcd quickly, sincc disavior. If reproductive
; had not yet dewlop0 adulthood to obtain
1 termitc reproductivc
:>tection from exccs:,ivc
1est material al-;o wcre
l· n 85°' cthanol, ¡\'lúch
;amplc were ohtained.
¡ collector, type of haancl weather conditiom
¡'(l
~cimem in the Nationa!
:. [N:\!:\IH], ancl the
(, NY [A.VINHJ. Somc
1e Estación de Biología,
ative Zoology, Han-arel
d in the scanning elecNickle and Collins in
ield studies for a book
, Specimens to be pho: absolute ethanol and
to drying. Tbey 11-erc
1g techniqucs clescribecl
fastcned 11·ith Elmer's H
:al stub for used in the
i-coated, cxamined, and
idge Scanning E lec tron
how spccies differences.
eithcr bccame they wcrc
Nere rendered ¡vhilc im-
TER:\IITFs
oF
CHA:\IELA
91
Il!Cr,ed in 8.J(·¿ Clhano] a> pen-and-ink Jine dra¡\·ings \\'it!J the aid of a
(amera lucida.
~ carly all figurecl specimem 1\'cre collected in Jal isc o State, but a few
spccies, reponed in the Jiterature as occurring in this rcgion, were figured
from spccimens obtained from localities outsicle of this region [e.g.,
imagoes of Xeotcnnes joutcli (Banks) 1920 ancl Xasutitermes nigYiceps
, Haldcman) 1t\5:3, and soldiers of Hoplotr'nncs amplus [ (Light) 1933]. \\'e
romideretl that for most species ('oldiers and imagoes alike) it was important
to illustrate interspecific differences of heads, both in dorsal ancl lateral
viein. ?llost of the features figured hercin are rendered as clear conci'c
im:tge' for the first time, and wc hope that they will greatly enhance
']JCrie-, rccognition ancl stimulate a renewed interest in termite researc!J
in thi;, rcgion.
The 11amcs of thc termites inclndecl in this papcr agrce with lh:tgc
listed by R. L. Araujo (1977). Data on clistributions and clepositories of
·.pcrimem, especially type material, wcre cataloguecl by Snycler (19-N\.
SPYC:IES OF TLR:\.f!TJ:S ÜCCLRRI:\G 1:\ THE CHA:\!ELA ARI:A
KALOTrR:I!ITlDAE
.\'cutcrmes jouteli (Banks) 1920
lncisitermes
nigritus (Snyrler) JCH6
lllluginiJiwnis (Latrcille) 1832
fJlaiYrr·j•I/IIIU.\ (Light) 1933
r IIICI".IOIIÍ ¡J.ight) 1933
.\Imginitnmcs hubbanli (Banks) 1920
Ca/raritennes
jJmr•inotus (Light) 1933
lougirollis (B3nks) 1918
r;, Yjl/olcrmn
{atulus ILight) 1933
!Jrcvis rWall:cr) 1853
RHL"\01 l R\ílTIP.\E
llcl!'r o! r 1 mn
alllfiH
Coj•lr•lnl!li'\ ilaS•lll
conn:xinolalns (Snydcr) 1~12·1
Snydcr 1922
FoLIA E:-;ro:-.wL. "IEx.
77
(I~ISS)
:\ICKL
Hoj,fulcnnn ruu¡Jlus Light 1'1:1~
,\1 iu urerotennc'
gracilis Light 1~~~~:~
sr:picnlriona/i, Light 19:13
\ledian ( \1) ve
R-,) aml cubit
1
ju't hcyoncl m
-lnzitclli!C'!
<TY}'iorlon I.ight l 1130b
1L'iledcri (Ik'ltcux) 190G
cmifer l.ight El~lllb
spccics 1, ma\ be
~pl'Cil'S 2
J){liVIlill\
\1
I.ight
.¡
iu/Jífonnans (Buckk\1 1863
¡;erplnu., (Banks¡ 1~1~0
s¡)l'cics 1
(~1111kr)
l ctding cclgc
EJ'2;\
(Snyder¡
<
<lo.sc to leadin
Light ¡q~~~)
11ígríccj;, (tbldcman) IR.'í;\
In ir iae
¡z., \\'ith :3-S br;
.,pccics)
R., with S-9 bra
_\'a\'lllilcnnc'
Jtlt'Yicanu.,
Tf'nllinntritnmr'\
3B) ....
1~1:1'2
( ;uulhamilr'rtlll'\
Fcnnn ¡wnan1ucnsí1
nmnmg m
1 Fig.
onh fe\\' -.hort
ex prc,,cd (Fig.
El:!~
l.c~tding
h.n
ro
THI·: TU{ \liTES
o!· nu:
,cp~11
CHA\II LA .\RIA
rttnnmg to lea•
dihlc \\'Cll dn
1 Fig. :) \)
1.\L\GOES
B;hc of [orcwing with :kí ,derotized Yeim, as countcd at suture
cdgc of wing '>cale. Fontancllc absent. Clypcus simple. Hcacl ami
pronotum .,parsely clothcd with sctac. Tibia! ;,puro, Z\:3:3 ...... -
,,
Ba<>c of fore\ving \\·ith onl) ~ sclcrotizcd YCino;, as countccl at suture
cdgc of \\·ing scale. Font;mclk prcscnt, though in somc specic~ in<ompicuom. Clypcus IJi.,cctcd or bilobed, though o.,omctime., IIl·
lOllo.,picuomh so. 1 Tcad ;md pronotum mt~;tllv hctYil' clothed \\·it h
Jll~lllY long ~tml o.,hort sct;tc. Tibi;tl spuro., not ;;:~l:~\
............ li
.\ntcrioi llLtrgtn of ;,ccoml m;trgin;d tootil of left nLmclible clongatcd. grcatn in lcngth th;m pmtcrim m~trgin of fiht marginal tooth
(hg. 2B. C:. F¡
... .....................
~;
cclgc '
atcd [rom
Forc\1· i ng sea le
Lipping it (Fi2
.t Jull com plcm
cxpandcd subsi
Forcwing -,c;tlc
dcarlv scparatt
m en tcd. .\ [andi
i
Hcad clongate<
Hcad subcircul
Fig·. 71F ..... .
.\ntrrior nLtrgiu of sccomlm;trginal tooth of ldt m;mdible suhcqual
in len¡;th to po;,tcrior margin of fir.,t marginal tooth (Fig. 2.\) .. :í
8
Foreti!Jia \Vith
38)
93
3
\ledi:m (\!) YCill uf forcwing nmning michray bet\H'Cil r:tdial 'lector
(R-,) :md cuhitw, (Cu) in pmxim;tl half ol \ring. lmt joining Rs
jmt hcyond midpoint (Fig. :lE) . . . . . . . . . . . . . .
CryjJtutenurs
\l numing midwa\ betwcen
1Fig. :lB) .. . . . . . . . . . .. . . . . .
j
R., :tiHl Cu to :qwx of
. . . . .. . .. .. .. . . .. .
forcwing
!
¡z., with ;;.:-; hr:mchcs (Fig. clH): l<lJo,;t] arolia pre.,em (cxccpt in
..................... ' ......... .
OltC
R, with il-~J hranchc-. (Fig. :lC); tar-.;tl arolia ahscnt .. Jfarginitcrmn
1
l c:tdi11g cdgc of forcwing 'cale o,traight. Hot!t R, ancl :\1 running
clmt· to lc:tding cdgc of forcwing. R-. without ln:tiH he, or \rith
on h few 'ihort branchcs. \Ltrginal tecth of ldt mand iblc \I"C<t kly
cxprc.,.,cd (Fig. :21J), lcss pointcd apically and 'iOtncwhat fuscd ....
...................................
Lc:tding cdge uf forcwing sct!c comcx, infLttcd. Rs more \l·idch
'<'p:tratcd frotn \1 (Fig. ~l.\). R, "·ith :l-~1 \l·c!J dcnclopcd hrandJc,
lllllllÍ11g lO lc;tding cdge of ÍOI"C\I'Íllg. ~Ltrgin;tJ ll'Ct !J of Jcft ITLIIIdilJ]c \1-ell dt·\clopcd, di-.tinn from each othcr, apict!Jy pointcd
'· Fig. :Z.\)
' ............................... .
Xcotr>nnc'
FLA ARL\
as counted at ~uture
eus simple. Hcad ami
Forcwing 'c:t!c comidcr;tl)ly longcr th;m hindwing .,cale. usualh ow'ILtpping it (Fi 0 . ~lll). \\'ing'i dc:tr. translun:nt. Lcft mandil>lc \l·ith
,t lull wmplcmem ol onc apical and thrce margin;t! tecth. ami an
cxpandl'Cl 'iub-.idi:tl) b:t'i;tl tooth (Fig. :ZF, G) ................. 7
1 spurs 3:3:3 ...... -
s, as countcd at 'iuture
gh in some spccic., inthough .,omctime-. in!y heavih clothcd \\"ith
3:3: ~\ . . . . . . . .
ti
left m;mdible cloug:tt·
of fjp,t marginal tooth
C: r1l ca rít t'l'lll n
Fon:\1·ing '>cale o11ly slightly. il at ;t!l, longcr th;tll hinch1·ing .,cale,
dcariy .,cparatcd from himlwing 'cale (Fig. ~11). \Ving'> oltcn pigmcmcd. ~\landihle-. not a-; above ............................. S
i
Hcad clong:ttcd iu dorsal \ÍC\1"
rig. :\G
................... "
(Fig. llA). \\'ing \ cnat ion as tn
......................................... llr'tcmlr'rmcs
,)
llead ,uhcirutlar in dm>;tl Yicw (Fig. llC). \\'ing 'cnation as in
Fig ..~1F
..............................
lcft mancliblc stthn¡u;tl
ll tooth (Fig. 2.\) ..
CojJtotcrmes
:1
8
Foretibia with ;) apical spurs ................................ 9
.'\ !CKU
FoLIA E;-.;ro:-.roL. Mtx. 77 (1 9HS)
01
Foretibia "·ith ~ apical spurs ............................... l:l
9
~lidtibia with
SOLDIERS
.'\asutiform spe<
ccaled bcneath
:1-± apical spurs. ::\Iandibles as in Fig. 2J ....... .
........... ... ..... .................... .......
,1ficrocerotermes
:\Iidtibia ,\·ith ~ apical spurs. :\fandiblc;, not a~ abo\·e ......... 10
lO
First and :,ccond marginal teeth of left mandible distinct, apically
pointecl (Fig. 2H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
.\ lanclibulatc sp
noth, cxtencling
')
First marginal tooth clistinct, apically pointed but posteriorly fused
with sccond marginal tooth, forming a broad, postcriorly roundecl
cutting shelf (Fig. 2L) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . l ~
ll
Posterior margin of apical tooth of lcft mandible equal in length
to anterior m~trgin of first marginal tooth. Right mandible witb
l\1"0 clistinct marginal tecth (Fig. ~H). Fontan:::llc indistiuct or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . AnuiJlotcnncs
Head capsule e
...... .
~ockcb
·'
Posterior margin of apical tooth of Icft mandiblc di:,tinctly Iongcr
than ~mtcrior margin of first marginal tooth. Right mandible with
only one wcll dcveloped marginal tooth, thc second tooth reduccd to
a wcak inflation on pmtcrior margin of first marginal tooth (Fig.
~I\.). }'ontanclle usually "·cll dewloped. slit-likc ........... Tennf's
1~
Hcad distinctly hroadcr than long. Fontanelle subcircular. ;\[anclibles
a-; in Fig. ~ (OJ. \Vings vcry long (> ~O mm) ... Tenuiro.1tritermes
cylinclric:
;¡pex; ncarly as
aspect (Fig. 151
-ocat te red seta e 11
scattcrecl short s
of ea eh tcrgum
t
long
brmn1
hcacl (3
(() blark
dolll in;tl tcrga 1
'>Ctac (Fig. l5B
"'
Total kngth witl1 1\·ing-; nol exccccling 13 mm. Hcacl rounded in
dorsal vic"·· Fontanellc usually wcll clcvelopecl. ~Iandibles as in Fig.
2I.. \\'ing \enation a-; in bg. 1E .................... /lmitermcs
Hcad :mhrinular or Iongcr titan broad. Foutanclle slit-Iikc. ;\Iandiblcs as in Fig. 2~. Wing' uot especial! y long ( < l:J mm) ....
................................................. 1Yasutitcrmcs
:'\ ~hll'i
:'\a<;u; lonical,
Font<mtilc absc
hehincl antcnna
Total length "·ith \\·ings greatcr than I:l mm. Hcad oval in dorsal
ncw. Fontanellc ,mall, oftcn olJ,,curccl. _\fancliblc'i as in Fig. ~:\I.
\\'ing venation as in Fig. 4B ................... Gnathamitennes
1'_.,·'
J n clor:,al Yicw,
1 Fig. l5C, F, I)
¡:ont:tnellc prcs<
,o me ;,pccics. ]
2:2:2 .........
-)
Head capsule ((
]J10' to an angle
tJ
VICW.
~\Ianclible'
capsule
••••
o.
o
Head ctpsulc el<
FoLIA E:-.:TO:'>IOL.
"IEx. 77 (1988)
:\!CK
H<>jilutennn rnllf!llll.
Light El:rl
\lcllian pi)
Miuocewtennn
gracilis Light
L~)3:~
sep!cntriorwli'
Light 19:\3
1
IR-,) :md cuh
ju'-L hcyond 1
~1 l1l i te rnu:s
.\1
cryptudon Light l~l30u
1L'izaleri (Dc'ltcux) 1906
t1L1ifer Light 19:lOb
spccics l. lila\ ue j){liVIlflll Ligh t 1~¡;¡~
~pc.:cics
running
1
!Fig·. 3B) ...
¡z, 1\'ith ;p; b
()
'))l'Cic>) .....
( ;natluun itcntlt''
tul!ifonllons (Burklc\ 1 18fi3
jJerpln:w
(l\anks¡ El~O
spccics 1
Fcn11n jlon!lllli/!'1/SÍI
l .c:tding cdgc
clmc to le:tdi
onh fcw -,hm
(~nnln) 19~:;
Sasutil<'nl/1'\
Jtlt'\<ÍcantL\
Lighl
1~~~~;¡
uig1 ircf" (J!a\dcman) IR.'í:l
1"1· y
TO THJ·:
TER\IITL~
t
Ol·
l.cading cdge
,eparatecl frm
THE CHA\IFLA ARI:.A
running to le
diblc wcll de
, Fig. ~.-\) ...
1.\IAGOES
Ba>c of iorc1ring with :\-:) ~clerot ized Yeim, a' counted at 5uturc
cdgc of wing scalc. Fontanelle ahsent. Clypcus simple. Head and
pronotum -,par,ely clothed with -.etac. Tibia! -.pur' ¿):3:3 ...... -
•i
Rase of forc,1·ing 11·ith only 2 sclcroti7ccl vcim. as countecl at suture
nlgc of 11·ing scalc. Font;mcllc prcscnt, though in sorne species incompicuom. Clypcus bisected or bilohed, though -,omctirne'i Íll·
<on-;picuou,Iy -,o. 1-Icad ;md pronotum usu:t!ly hcaYily clothed with
m:my long ;md ,Jwrt seLtC. '1 ibial 'Jllll'., not :\:3:~\ ............ ti
Anterior nt:trgin of second m:uginal tooth o[ lcft m;mcliblc clongated. grcatcr in lcngth th;m pmtcrim m:ugin of fir-,t marginal tooth
(l··ig. 2!~. (:. E) ............................................. ~~
x )JlC))ecl (Fi¡
Forcwing scah
Ltpping it (F
:t full com ple1
cxp:mdcd sub
Forcwing sctJ,
clcarly separa
lllCII te el. .\ {all<
7
Hcad clongat
lig. 3G ....
Hcacl subcirct
Fig. ~1F .....
.\ntcrior m:trgin of sccond marginal tootlt of ldt mandible sulwcptal
in lengtlt to posterior margin of first m;trginal tooth (Fig. 2.\) .. :1
8
Foretibi a 11·itl
.:\ICKLF
3
LT CoLIJ:\s.
TEK\IJTES
oF
CHA.\IEL\
93
\Iedían ( \1) v-e in of lorewíng nmning midway betwcen radial '>Cctor
iR'>) ;md cubítw, (Cu) in proxím;¡] half ol wing. but joining Rs
ju-;t hcyond midpoínt (Fig. ~H-) .... _ . . . . . . . . . . . . . CryjJtotenurs
.\1 nmníng midwav between R., and Cu to apcx of forewing
(Fig. 3B) ... __ ........... _. . . . . . . . . . . . . . . . . . . . . . .
!
l
Rs \\'Íth 3-N IJr:mches (Fíg.
:m):
tar.sal arolía prcseJJL (exccpt in onc
...........................................
Rs with N-~) branchc' (Fig. :\C); t:lr'>;ll arolia aiJsenr .. .\Iarginitennn
.J
Le:1ding edge of forcwíng 'cale ..rraíght. Botlt ¡z, and ;\1 running
clmc to le:1cling cdgc of forewíng. ¡z, without br;mcl1e, or with
onh few o,l10n branches. .\ Ln ginal teet h of Id t. mandiblc vrea kly
npres'>ctl (Fig. :20), less pointccl apirally and '>Oillewhat fused ....
Calcaritcnnn
Lc:1ding edgc of forewing selle comex, inflatcd. Rs more \\'idcly
'cp;uatcd from .\1 (Fig. ;\.\). ¡z, "·itlt :l-9 V\'Cil dcn~lopcd hranchc,
runníng to leading cdge of lm-cv1·ing. Margina! tccth o[ !eft m:l!ldiblc wcll dnclopcd, di'itinn from cach othcr, apícally pointed
¡Fig. :¿_\)
...................................... ' ..
'
1s counted at suture
1s simple. Head and
FoJT\\·ing- .,tale comidcrably longer th:m himhring -,cale, u:,u;dh ovcrLtpping it (Fig. ~lll). \Ving-; dc:11. transltHcnt. Left mandiblc v1·ith
:1 full tomplcment of one apical :md tln-ce margin:ll teeth, aml an
cxpandcd subsidi:11y ba.~al tooth (Fig. 2F, G) ... _ ... _ .... _ .. _. 7
spurs 3:3:3 . - .... -
as counted at suture
in some specie~ inwugh sometime-; inheaYilv- dothed vvith
1
Forc:\1·ing st:lle ouly :.lightly. ii at :di. longcr than hindv1·ing '>cale,
dcarly >Cparatcd from hímh1· ing sede (Fig. ~¡ l ). Wíng'> oitcn pigmcntccl. .\l:twliblcs not as abovc . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
:3: ~l ... ' . . . . . . . . . ti
eft mandiblc elong:\l' fírst marginal tooth
e>
........ ' ...... ' .
i
IIcad clongatcd in dmsal \IC:\1" (Fig. !lA). \Ving Yenatíon as in
ri¡.;. :lG ......................... _ ....... _....... Hetemtcrmes
,)
Hcad .,ubcircubr in clors;d v-iew (Fig. llC). \Ving vcnation as in
Fig. :1F ....... _....................... _.....
CofJtotermes
:ft mandible subcc¡ual
tooth (Fig. 2.\) .. 0
S
Forctibia with 3 apical spurs _ ................ .
......... 9
01
FoLIA
E:'>TO:\IOL.
Foretibia "'ith ~ apical spur' _ .. , .. __ ..................... _ _ lJ
9
:\IC:KI
.\le..:. 77 (l9RS)
SOLDIERS
.\'a~ut
.\lidtibia with :1-1 apical spurs. i\Iandibles as in Fig. 2J .. ___ ...
.............................................. ¡\IiCTocerotermes
.\Iidtibia "'ith ~ apical spurs . .\fandiblec, not as ahü\e _. _...... 10
10
First and ~econd marginal teeth of left mandible distinc:t, apically
pointecl (Fig. ~H) _.. ___ ......... _ . . . . . . . . . . . . . . . . . . . . . . . . . . 11
.\ Ianclibulate s
uou,, extcnclin
C)
First marginal tooth clistinct, apically pointecl but pmteriorly fused
with seconcl marginal tooth, forming a broad, po-.teriorly rounclecl
cutting shelf (Fig. ~L) ... _........ _... __ ... __ . . . . . . . . . . . . . . . 1~
ll
Posterior marg!n of apical tooth of left mandible equal in length
to anterior m~trgin of first marginal tooth. Right mandible with
two clistinct marginal tceth (Fig. 2H). Fontanelle indistinct or absent ...... _ .. _..... __ ........ _................ ,. Anoj)/olcrmcs
Posterior margin of apical tooth of left manclible di:otinctly longer
than :mterior margin of first marginal tooth. Right manclible with
only one well cleYeloped marginal tooth, the second tooth recluced to
a weak inflation on pmtcrior margin of first marginal tooth (Fig.
21\.). Fontanelle usually well deye]oped, slit-likc ........... Tennr>s
Tot:.Jl lcngth \\'ith wing,; uot exceccling 13 mm. Head rounded in
dorsal yie11·. Fomanellc usually well ckvelopecl. Mandibles as in Fig.
~L. \\'ing· \Cnation a<; in Fig. 1E .......... _....... ,. "'1mitermcs
Total length 11·ith \rings grcater tllan 13 mm. Head oyal in dorsal
\"Íew. Font:mclle 'mall. often o!J,curecl. 0Lmcliblc'i as in Fig. 2:\I.
\\'ing yenation as i11 fig. 4R
.. _......... Gnathamitcnne1
Hcad suhcinular or longcr than lJroad. Folltanelle slit-likc. Mandihles as in Fig. 2:.\'. Wing, not cspccially long ( < Ei mm) ....
................................................. 1Yasutitcrmcs
Heacl clistinctly hroader than long. Fontanelle subcircular. :\Iandibles
a-; in Fig. 2 (0). \\"ings yery long (> ~O mm) ... Tenuirostritermcs
ifonn sp1
<ealecl beneath
In dorsal YÍC\1
1 Fig. 15C, F, I
Head capsule
-;ockeb . _....
•
.:\a,u-; cylinclri
:tpex; nearly a~
aspect (Fig. l.
'cattcrcd ~etae
scattercd short
of each tcrgun
conical,
long ;l'; hcad e
brmrn tu blad
dmn in:tl tcrga
'etac ( Fig. 1:)
'\;¡<;u;
hmtaaclle abs
bchind antcnn;
i:ontanclle pre
,omc opccics.
Heacl capsule 1
ing to an angl<
Yiew. ~Iandibl1
capsule ..... .
Heacl etpmle e
:'\rc:KLF IT CoLLL\s.
o
o
o
••
o
•••
o.
o
•••
13
o
m Fig. 2J ....... .
j\f icroce ro termes
; aboye . . . . . . . . . 10
9:)
SOLDIERS
:'\asutitmm spccies: mandiblcs inconspicuous, grcat1y reduced, conccaled bencath labrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ~
.\lamlibulate specics: mancliblcs enlargcd aud morc-or-lcss conspiccxtending 1\Tll hcyond margins of labrmn . . . . . . . . . . . . . . . l
UOLh,
ible distinct, apically
11
'
TFR~llTES OF CHAMEL\
o
•••••
o
•
•
•
•
•
•
•
•
•
but posteriorly fused
, posteriorly roundccl
1:2
••••••••••••••••
o
lible equal in length
~ight mandiblc with
.elle indistinct or ab.. . . . . Ano,blutenncs
:ible distinctly longer
Right mandible with
:ond tooth reduced to
marginal tooth (Fig.
: ........... TennPs
m. Head roundecl in
~Ianclibles as in Fig .
. . . . . . . . . Amitenncs
Head onl in dorsal
liblcs as in Fig. 2.\I.
.... Gnatharnitcnnei
~anelle slit-like. ~lan­
rlg (< 15 mm) ....
. . . . . . Nasutitermes
ubcircular. .\Iandible~
... Tenuirostritcrmcs
In clon,al \'JCW, hcad cap'>ule comtrictecl behind antcnnal sockets
IFig. l5C, F, l)
Tenuirostritcrmes briccae
Heacl ctpsule oya] or pyriform, not constricted behind antenn:tl
o;()CI,et~ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Nasu-; cylinclrical, clongate, awl nearly as narrow at base as near
:tpex; ncarly as long as head ctpmle behincl nasus as scen in lateral
:t'pect (Fig. l5G). Head capo,u]e amhcr colorccl, witlt only a fe\1·
.,cattcrcd 'cüe (Fig. l:"íD) .. \bdominal terga ncarly bare, with \\'Íclcly
sctttercd short sctae ancl only Z-3 large set:te on the posterior margin
of eacll tergum (Fig. 15A) ............. Xasutitermcs mexicanus
:\a<;th conictl, broader at base than at apcx, only about one-half as
long as hcad capsule bchincl na'm (Fig. 15H). Heacl Gtpmle dark
br01n1 lo bl:tck in color, wíth numcrous long setae (Fig-. FJE). AlJdom in;tl tcrg;t 11· it h numcrous short ancl long, raudomly scauered
'ctac (Fig. EíB) ........................ Xasutitcrmcs nigricc/J\
Fonta:wilc ab,erlt. Eyes usually prescnt, locatccl on siclcs of hc:1d
hehincl antcnnal sockct. Tibia] s¡mrs ~l::l:3 .................... :J
Fontanellc prcsent, tlwngh inconspicnous or clifficult to locate in
:,omc ;,pccic,. Eyes absent. Tibia! spurs eithcr 3:2:2, 3>1:2, or
{) • C) • ')
• • • • . • , • • • • . , • • • • . • , . , , •• , , , , , , , , • , , . • • , , , , • • , • • • • . • • ,
1-1
He a el ctpsule compresscd, distinct ly phragmotic, with forehcacl rising to an angle o[ nearly 00° with the horizontal as sccn in lateral
view. ~\Lmdibles reduccd in size to !css than 0.3x lcngth of head
capsule ..................................................... 6
Heacl ctpmle elong;ltccl; forcheacl in lateral view sloping at an angle
.:\'ICK
FouA E:--;To,IOL. \hx. íí
( 1~)~~)
ll
of 60'' or le.,, with horizontal. \IamlílJlc., "·el! dc\'eloped. cxtcnding
in fro11l o( hcad, at le<t'il 0.3x length of hcad . . . . . . . . . .
. lJ
Lateral lengt!J
less than l.i n
length of arti
Frontal ridgc of ltcad ctpsulc crenelate, cntirc. not medially enLtrginated (Fig. 71)) .. \ll "purs oll foretibia nearly cqual in ~ize ... 1
Front:tl ridge of head capsuk smooth or 11·cakly punnatc with a
medial. deep. ll-shapcd emargination (Fig. 5F). Foretihi:t with onc
grcatly enlargccl apical s¡mr and two smaller emes . . . . . ..... 8
~~
Sides of head
(Fig. 5D). Se<
distinct cusp;
6E) ....... .
Frontal ridgc of hcad fineh < reueLttn1, without a dcep depre"ion
IJchind ridge. From abO\e, head suhquadrate, not significmtly con:otriued hchiml fronLtl ridgc (Fig. 7D, }') .... CryjJiofi'IJJII'\ fatulu'
ln dOJ-,;t! Yiew, head subquadr;ttc ........ Ca/c!lrill'l"llli:l /ongicollis
(j
11-
.\ntcrior margin of pronotum cmic:tvc (Fig. 7.\). but neitlicr inci,('(l no1 \l·itlt a \'-shaped emargitLttion. \Iandihle dentition a~ in
Fig. li.\ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sr·otcrJ/11'1 joutl'!i
.-\ntcrior margin of pronotmn medially inci-,cd with :1 decp \'-shapcd
cmargination (Fig. !í,\). :-randible dentition differing from ~thcnc
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lO
Sides
(Fig-.
tinct
(Fig.
of hea'
5C). se,
cusps; r
6F) ...
Inner
a fcw
tibia!
dially
margin
serratic
spurs 3
notche
.\Iandibk~
w
Tibia! spurs
shaped, with
l!í
Thinl :lllteunal :tnicle greatly cnlargcd, ;2: :trtidc-, ·l-~l wmbincd.
.\ distinct raised ridgc bctwcen from. aml ,-cncx, thick.cst abO\c.
:-I~mdib!e, a' in Fig. hG ................. Jlarginitcnnc.1 lwúlJard,
Thircl antennal anide < articlc' ·!-7 combiued. Ridgc not clistincl
:1lthough m:11 be tltick.cncd ahO\e antcnnal o,ock.ct~ . . . . . . . .
ll
Lateral lengtl
6D ....... .
Lateral Jengt
aboYe ..... .
FronLtl ridgc of head dceply rugosc, with :t deep deprc~sion beltiml
ridgc. From :1 bo\"C, lte:td constrictcd hch iml front<tl ridgc (Fig.
8.\. B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . CryjJtutr·nncs lncun
Ju dors~d ' ic"', !tead rectangula te, d ist i nct l y longer th:tn \\.id e
\Fig. :"JF) .............................. Calcaritcrlllt'.l parvinotu5
Lateral lengtr
as articles 4-í
Head pyrifor
ed, directed
Hcad rectan
reduced, ap¡;
dorsal regio!
](j
Head with a
97
~ICKU~ ET CoLLI:\S. TER:\IITES UF CHA:\1ELA
8)
11
l dcveloped, extencling
l ................. q
Lateral length of hcad not mually exceecling 2.5 mm and gencrally
less than 1. 7 mm. Thircl antcnnal article as long as but not exceeding
length of articles ·1-5 combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
re, not meclially cmar-1
dy equal in size
eakly punctatc 1rith ;1
F). Foretihia with onc
r ones ............ S
.lccp deprcs~ion bclliml
el frontal ridgc \ Fig.
.. Cryptutcrl/lcs lnn!i1
12
Lateral length of head > 2.0 mm. J\fandible dentition as in Fig.
6D . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lncisitermes nigritus
Lateral lcngth of head < 2.0 mm. ~Iandible dentition not as
abovc ..................................................... 13
13
tout a dccp dcpl-c,.,ion
not significantly com. Cryptutcmu:1 faltfiu,
Sides of ltead weakly com·ex in dorsal view; head postcriorly rounded
(Fig. 5D). Second marginal tooth on left mandible with only one
distinct cusp: right mandible robust, apically strongly recurved (Fig.
6E) .................................. lncisitcnnes platycephalus
Si eles
(Fig.
tinct
(Fig.
:alrariterllles /ongicullis
:tly longer than 11·ide
alcaritermes pan~inotu5
Lateral length of head > 3.0 mm. Third antennal articlc as long
as articles 4-7 combinecl .............. lncisitermes marginipennis
H
~- 7A), but neither inmdiblc dentition a~ in
.... Xeotcrmes joute/i
Inner
a few
tibia!
clially
of hcad parallcl in clorsa 1 Yiew; head posteriorly
5C). Seconcl marginal tooth of left mandible with
cusps; right mancliblc more slender and apically less
fJF) ................................. Jncisitermes
trunca te
two disrccurved
emersoni
margins of both manclibles smooth, lacking teeth, cxccpt for
scrrations at base of each mandible and concealecl by labrum;
-;purs 3:2:~. Pronotum flat with anterior margin entire or menotched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . l.~
~lancliblc-; with teeth or tooth-Iike moclifications on inner margin.
Tibia! ~purs mually 2:2:2 but may be 3:4:2. Pronotum saclclleshapcd, with a broadly rounclcd upcun-cd anterior lobe . . . . . . 16
el with a cleep V-shapcd
differing from ~dJo\c
1
........... _......
lO
15
articles ·1-~J combincd.
vertex, thickcst a bmc.
\Iarginitennc,\ llllbúard,
ned. Riclge not clistinct
;ockcts . . . . . . . . . . . . 11
Head pyriform in dorsal \ iew (Fig. lOA). Fontanelle greatly enlarged, directed antcriorly (Fig. JOB) ............ Coptotenncs crassus
Hcad rectangulatc in dorsal view (Fig. !OC). Fontanelle greatly
rcduced, appcaring as a pinholc between two inflations on anteradorsal region of head (Fig. lOD, E) ........................ .
. . . . . . . . . . . . . . . . . . . . . Hcterutcrmcs uurr:us cmm,cxinotatus
JG
IIcad "·ith anterior triangulatc cone (Fig. 1 lB, C).
~Ianclibles
grcJ.t
FoLIA
!JS
17
E:-.:TO;\IOL
\IEx. 77 (l9SH)
NICKL
h clongated, narrow asymrnetriral and curvccl downward m lateral
Yicw (Fig. 1 L\, C) ........................ Ter111cs fJanamacnsis
Head more ova
bilobed ..... .
""ithout the ahove combination of character-, . . . . . . . . . . . . . . . . 17
\landibles gre<
invurvcd along
ded cutting slH
labrurn (Fig. 1
lnner margim of mandí bies
~erra ted
a long the ir cntire length . . l S
lnner margim ol mandiblcs each with a single tooth . . . . . . . . . El
18
In dorsal view. head oval (Fig. 1:!A); mandih]e, finely serratecl.
their length > 0.:) times length of heatl cap-,tile (Fig. lZB) ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ,\I irruccrutcrmcs gracilis
,\/audibles less
disti nctly i ncur
l nner margin o
ing as preapica
llead rectangulate (Fig. l:!C); mancliblc, mar:-.ely serratccl, their
length < O.:J time-, lcngth of hcacl capsule (Fig. l:!D) .......... .
¡\[ icror!Totenncs scptentriunalis
1nner margin o
ed looth ....
Mandibles shon, hasally stout; head rectangulate, median suture
well developecl (Fig. 12E). Facial profile with a high angled ridge
(Fig. 12F). . . . . . . . . . . . . . . . . . . . . . . . . . .
HujJlotermes arnplus
Lateral length
\'Cioped medial
pem \\·eakly o
t\I andibles elong;tted, nriously toothcd hut general! y slender in appear:mce, either straight or sickle-shaped (Figs. 13, llD, E, F);
head oval or rectangulate; median suture inconspicuom. Facial profile 'iloping anteriorly. without a high ridge .................. 20
Lateral length
median tooth
C:lypcus cli-,tinc
\[audibles gcneralh, straight. ;p; long as or longer th;m lateral length
................................................... 21
of head
i\fawlihles generally recun·ecl in shape, slwner tlt;lll later;t! lcngth
of hcacl (:t!though 'itraight in one sperie-;) ................... 23
,\fandihles with a shehT<l cutting eclge along di-.tal half of inner
nurgin (Fig. liD) ................. GnatluunifrTillr\ tu/¡ifnnnum
\i;mdibles 11'it!tont a cutting shclf, but bearing a 11·cll dcwloped
ll!ar¡J,in:tl tooth near ba'ial thinl of inner mandihubr margin .... :!~
llcad tapcring :mteriorly: mandiblc as in Fig. HF; clypcal margin
not distincth hilobed ................. Gnathamitermes perplexus
'\EOTERMES Holu
, toul Icngth usuall)
hcHh (in dorsal vie1
llwy ;tre clistinguisl
1 ;tctcrs: ( 1) anterior
;tlld 1~) third anter
1 ombincd. Jmagoes ,
o( wing vcnation ar
i' (_haracleristically
w the Cu. Species <
mental moisture sup
:Collins. l%9). Onl)
1\' ICKLI:
downward in lateral
Termes panamacnsis
................ 17
~ir
entire length . . 18
le tooth ......... 19
iihle~
finely serratecl.
>sule (Fig. l:!B) ....
áocerotermcs graci li.1
1ar~ely serratecl. their
'ig. l:!D) .......... .
termes scptentrionalis
ulate, median suture
a high anglecl ridge
HofJlotermes am fJlus
nerally slender in apígs. 13, llD, E, F);
nspicuous. Facial pro.' ......... ' ..... 20
ET CoLLI'\s. TJ<:R.\IITJ<:s OF
CHA:\IELA
99
Head more oval; mandibles as in Fig. 11F: c!ypeal margin distinctly
bilobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gnathamitermes species
Mandibles greater than 0.8x lateral length of head, only weakl~
invurved a long their length. 7\I arginal tooth cleveloped as an extended cutting shelf with a deep notch at base of mandible, beneath
labrum (Fig. 13C, D) ..................... Amitermes cryptodon
,\Iandibles less than O.Gx lateral Iengt!J of head, moderately to
distinctly incunnl .......................................... 2±
lnner margiu of each mancliblc anned with a cutting shelf terminating a-; preapical hook (Fig. l3E, F) .......... Amitcrmcs species :!
hmcr margin of cach mandible armed with a distinct apícally rounclecl ·toot!J . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2:!
Lateral length ol hcad > 1.25 mm. Manclibles mbust, with well dcn:loped median teetlt ;tri'iing near midpoint of inner margin. Clypcm weakly or indistinctly bilobecl (Fig. l:lG, H) ........... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amitermcs wheclcri
Lateral length oi head < 1.25 nun. ~1 audibles with smaller loba te
median tooth arising somewhat more toward base (Fig. 13E, F).
Clypeus <listinctly hilohed ................... Amiternu:s specíes 1
er than lateral length
................ 21
r tii;m lateral length
.... ' .. '.' ' .. ' ... 23
distal half of inner
ni termes t ubiforma/1.1
ng a 11·ell cleveloped
bular margin
1'1E; clypeal rnargin
1amitcrmes pcrplc:ws
KALOTER\l!TID.u::
~EOTERMES
Dry-wood Termites
Hohllgrcn 1911. Solcliers of Seotr:rrncs are generally large
> 8 mm), with elongated mandibles, rectangulate
hc;td> (in dorsal vie1\'), aJI(] 11·ell developecl, somctime'i pigmented eyespots.
I'llC'y are distinguished from lnrisitermr'.\' soldiers by the following chal ;tcters: ( 1) anterior margin of pmnotum concave but never cleeply incised.
;tnd (..!) thircl antemul article mually not as long· as artícles 4 ancl 5
tombined. Imagoes of ll1i'i genus are most easily recognized on the basis
()f wing venation aml rnandible dentition. The M vein of the forewing
i-, characteristicdly sclerotized aml situated closer to the Rs than it is
10 the Cu. Specics of this genus are dependent upon a higher enviranmental moisture supply tl1an species of cither Jncisitenncs or Cryptotermcs
~Collins, 1%9). Only one species is li'>tcd from the Charnela area.
1tot:d length usually
lOO
FoLIA ENTO:\IOL.
:\[Ex. 77 (1988)
NICI
~DOD
úD
e
July
19 and Augus
branches or roots o
~pecies has a high n
tures unless the wo1
in the roof). The
Florida, Bahamas,
and on the weste
J<tlisco (see be lo>'
\ibling species cor
liiÍncd: 13 soldien
west beach of El I
.\ISCollins) VI-29-l
Chamela. 5 soldie
l ~IH-1 [NMNHJ; sa1
living tree. 2 soldi
I~CISITERMES
merous species sh
imago with anteric
pmterior margin e
midway between l
of thc soldicr de<
the third article ~
articlcs. Four spec
~trea.
Fig_ 2. Imago¡nymph¡'worker man<lihles of genera of tcnnites in thc
mcla, Mexico. A. Ncotcrmes; B. Incisitcrmes; C. A1arginitermes; D.
CryfJlotermcs; F. Hclerotcrmes; G. CojJtotenncs; H. AnojJ/otermes;
T- Jficrocerulermcs; K. Termes; L. Amitennes; .M. Gnathamitermes;
O. Tenuirostritcrmes.
vicinity of ChaCalr-nritermes; E.
I. HojJ/otf'rmo;
1\'. Xasutitarne<;
N eotermes joute1i (Banks) 19~0. This termite is casi! y rccognized by the
large sizc of the solcliers (lateral lcngth of hcacl 2.1-2.8 mm for shorthcadcd soldicrs ancl 3.()-~).4 mm for long-hcadecl solclicrs), the conGtve but
noL incisccl anterior margin of the pronotum, ancl small, clarkly pigmcntcd
cycs (Fig. 5E, F) . . \late-; are golclcn to light hrown in color, with a
total body length of 8-~J mm. Head lcngth avcragcs 1.5 mm ancl hcad
width 1.7 mm (Fig. 9E, F). \Ving lcngth is about 11 mm, wiclth about 3.0
mm . . \ colony collcctcd ;lt Barra de I\' aviclacl procluced al ates between
I ncisitcrmes emer
onlv from the solcl
of the heacl ranges
!tctd is distinctly
pmterior margin
lcft mandible ha'
ouly one, and tb
apically (Figs. 6E
from "thc sound
\rorkcrs of two ~
;n ailable, since tb
containing identif
is obtainccl, we 51
the date of matm
ancl prcserving, b
NICKLL ET CoLLINS. TER'\IITES oF CnAMELA
OD
e
:JQ
JD,
JQ
JU
in the
les; D.
!termes;
termes:
vicinity of Ch:tCalraritermes; F..
I. Hof!lotermc';
;\/ . .Vasutitrrme1:
ily rccognized by the
~.l-2.8 mm for shortiers), the concave but
al!, darkly pigmented
wn in color, ,.,·ith a
~s 1.5 mm and hcad
mm, width about 3.0
luced alates between
101
July 1!) and August 2, 1968 (MSC, unpubl.). Colonie> are found in dead
branches or roots of living trees, particularly at lower elevations. Since this
~pecies has a high moisture demand, it is unlikely to attack man-made structures unless the wood were repeatedly rendered moist (e.g., because of leab
in the roof). The distribution of this species is disjunct, occurring in
Florida, Bahamas, Cuba, Trinidad, Tobago, eastern Mexico (Veracruz).
and on the western coast of l\Jexico in Sinaloa (near Mazatlan) and
Jalisco (see bclow). Further studies m ay reveal the occurrence of a
~ibling species complex (see Nickle and Collins 1989). Specimens Exalllincd: 13 soldiers, 127 nymphs. l\IEXICO: falisco, Barra de I'.'avidad,
\\'est beach of El Dorado Hotel. '1 soldiers, 31 nymphs (Col! DANickle &:
_\fSCollins) VI-29-1981 [NMNH]; falisco, Estación de Biología, UNA.\1,
Chamela. 5 soldiers, 50 nymphs (Coll. DAN ickle & MSCollins) VI-29l 9H.f [NMNH]; same data. 2 soldiers, 30 nymphs [NMNH]; same data. In
liYing tree. 2 so1diers, lti nymphs [N-'INHJ.
I~CISITERMES
Krishna 1961. This genus was erected to include numerous species sharing the following characters: (l) 1eft mandible of
imago with anterior margin of second marginal tooth slightly longer than
posterior margin of first tooth; (2) l\I o[ forewing unsclerotized, running
midway between Rs and Cu; ami (3) anterior margin of the pronotum
of the soldier cleeply incised or emarginated, the femora swollen, and
the third article enlargcd and more heavily sclerotized than subsequent
articles. Four species of Jncisitenncs havc been collectcd in the Charnela
are a.
Incisitennes emcrsoni (Light) 1q33. Inci.1ilennes cmcrsoni is known
onlv from the soldier caste. Soldiers are small for the genus; lateral length
of the head ranges from l.!JH to Z..H mm. In dorsal view, the shape of the
hc;td is distinctly rectangulate, with siclcs of the head parallel and the
po-.;terior margin truncate (Fig. SD). The second marginal tooth of the
lclt mandibJe has two cusps, unJike that of platycephalus, which has
only one, and the righ t mandible is more slemler and less recurved
apically (Figs. 6E, F). Light (1933) collected this termite near Colima,
hom "thc souml wood of an old, much-decayed, harclwood stump, with
\l"orkers of two genera uf subterrancan tcrmite,." 1'\o flight elata are
a\ ailable, since the ala te has 11ot yct been describccl. If a picce of >mod
containing idcntifiable !. ernersoni soldiers and nymphs or pseudergates
is obtainecl, we sugge-;t that cfforts be made to rear ;dates and record
the date of maturation ..\lates taken at lights are always worth keeping
ancl presen-ing, but dcscriptions of the ;date of this species should be
102
FouA ENTO:'.lOL.
;\lEx. 77 ( 1988)
:\'tCK
anides; (l) :\1 yeiJ
(1) wings paJe, \CÍI
Fig. 3. Imago "ings of termitcs of the families Kalotcrmit i<lac and Rhinotermitidae i11
the vicinity of Charnela, Mexico; :\ . .\',·otcnnes; B. lnci.IÍ/ennn: C. .\lar¡;initcrllll''·
D. Calrm·itnme.,; E. CryfJtotcnnes; F. Coptotcrmn: (;. llctnotennn. 1\1
median ICÍIJ
o! folT\\·ing.
=
Scyeral features '
with the monotypi
othcr membep; of
tcrs, sharecl with J.
third antennal arti<
than next four arti
cially raisccl oYer a
margins stra ight ale
tooth of the left n
ginal tooth, but "
trailiug to third m
rising bladc to a ¡:
ly grcater in lengt
1cj. Figs. GB ancl t
doscr to midpoint
terior thinl of pr
tlcep, \'--,hapcd mt:
arolia are absent i
¡t hough present in
hiological affinitie
testinal protozoa
divergens Kirby, a
form the affinities
gen) 1~l:">R, share'i
tionship with thes<
Incisitenne1 111a1
or hu ilclings, or in
scr\'ations by Roge
Rogers reportee! ti
rainy SC<bOil, COllll
t hrough June. Th
extensiw burning
construction. !neis
in the states of 0
aho been collecte<
1
made only on fonns associatc:d \rith -,oldic:rs. This species
from type material collcctcd in Colima.
1s
known onh
Incisitermcs marginipcnnis (Latreille) I8:l2. Solcliers of Incisitermes 1/larginipcnrzis are the largest for this genus in tbc area. Lateral Iength of the
l1ead ranges from 2.19 to 3.31 mm (Light 1933). As with severa! other
'pecies of this genus, therc are two forms of soldiers: long-headecl aml
short-headccl (Fig. :í,\, B). The primal} charactcrs usecl to distinguish this
<pecies are ( 1) its largc third amen na] article, which, though not as large
a'i that of 1\1arginitenncs hubhardi,. cquals or exceeds the combined lengths
of articles ·1-7; (2) its o\'erall large sizc which is greater than 10.0 mm;
ancl (3) thc deeply incised anterior margin of the pronotum. Imagoes are
recognized using the following characters: ( 1) head yellow-brown, Ionger
than wide, posteriorly rounded in dorsal view. with wiclcly scattercd
'ihort !>etae: head length 2.20 mm, width 1.80 mm; (.!) antennae "·ith li'
Incisitermcs nigrit
are larger than e
:\JcKu
D
=:::
__
z=--~-~~.
FT
CoLu;-;s.
TER\IITES
oF
CIL\\IELA
103
articles; (3) .\l Yein brancbed subapically, equidistant from Rs and Cu;
('1) wings paJe, Yeim darker; wing length 16.0 width '1.5 mm.
====o
__
-.::-=-
~......-.-..-~
~~~0)
E
~---~~
G
====~
~!
ami Rhinotermitidae in
·mn: C: . .\lnrginitallll'':
nnes. 1\I
)Ccies
IS
= median
1cin
known onlv
of lncisitermes 1narLateral length of thc
s with severa! other
TS: long-headed ancl
~d to distinguish this
though not as largc
:he combined lengths
:ater than 10.0 mm;
motum. Imagoes are
yellow-brown, longer
ith widely 'cattered
2) antennae with l t'
SeYeral features of lncisitermes rnarginipennis suggest closer affinities
,\-ith the monotypir -,pecies Marginitermes hubbardi than it has with
other mcmber<> of the genus lncisitenncs. The following soldicr characters, sbared with JI. hubbardi, scparatc it from lncisitermes spccies: (1)
third antennal ;trticle relatively more heavily sclerotizcd, greater in length
than next four ;trticle'i comhincd; (2) frontal ridgc well developed, especially rai,ecl m er ;mtennal 'ockets; (3) manclibles elongated, their lateral
margim straight ;tlong most of distal two-thirds of Iength; second marginal
tooth of the left mamlible with primary cmp arising closc to first marginal tooth, hut with a long, extended cutting edge or secondary cusp
trailing· to thinl marginal tootb, \\'hirh in turn commences as a gradually
rising bladc to a peak and situated near thc miclpoint of a ridgc slightly greater in length than thc total length of the second marginal tooth
(ej. Figs. 6B ancl 6G); right mandihle with first marginal tooth arising
closer to midpoint o[ inner margin of mandible than to apcx and (4) anterior third of pronotum upcuncd; anterior margin serrated, with a
deep, V-,haped mcdi~1 n cmargination. Furthcrmore, in the imago, tarsal
aralia are abscnt i11 both l. marginipenni1 and J1m·ginitermes hubbardi
(though jJrC'ient in other spccics of Jncisitermes). Light (1933) also stresscd
biological affinitic.-. of l. marginipennis with M. hubbardi: shared inte~tinal protozoa (Staurojoenina sp., ,\letadeveiCU1'Ína sp., Tricercornitus
divergcns Kirby, ancl Oxy11wnas sp.). Future studies may cventually conform the affinities of these two spccics (another spccies, l. minar (Hagen) l q:)t-1. sharcs sc\eral of thcse characters as 'rell, and its close relationship with the-,e two ~pccie'i may eventually be confirmed).
lncisitcnnc.l llínrginijJI'nnis is founcl in the souml dry wood of fences
or buildings, 01 in dcacl branches on live tlees. Light (1933) included ob.,crvations by Rogers on flights of l. mnrginipcnnis alates in Guadalajara.
Rogers reportecl that flights began severa! days after the beginning of the
rainy scac,on, cmnmcncing at niglttfall, starting in May, and continuing
through June. This tcrmite may be less ahundant than formely, due to
cxtcnsive burning of wood a-; fue! and increased use of treated timber for
construction. lnci,itermcs 111arginipennis is widely distributed in Mexico
in the states of ::'\ ayarit, Colima, Jalisco, ancl Distrito Federal, and has
aho bcen collcctcd in Guatemala.
Incisitermes nigritus (Sn;der) EJ16. Soldiers of this termite (Fig. 8 C, D)
are larger than emersoni ancl platyccphalus but smaller than margini-
104
FoLIA E:-.Tü:\IOL.
MEx. 77 (l988j
~~
~~~
hg. 4. !mago "·ings of rcpresenlati\e termitc spccics of thc Tcrmitidae in Lhe vicinilv
of Chamela, ,\Icxico; A. i\Iicrocerotermes: B. Gnathamitermcs; C. AnofJlolcrmes; D.
Termes; E. -~mitennes; F. Xnsutitermes; G. Tenuirostritenuc.•.
pennis. Besides size, however, thc hest means of characterizing thi-; species
is mancliblc dentition (Fig. 6D). This littlc-knmm tcrmite was first dcscribcd by Snyder ( EJ46) from a shipmcnt of wood from Guatemala to the
Unitccl States. It was first founcl in .\[cxico in 1069 bY J. Collins (son of
MSC) in tiny twigs on living trccs at Barra ele :\" aYidad (Collins, unpubl. obs.) ancl was later collected frcquemly in thc heartwood of ste1m
and branches of living hardwoocl trecs near the shorelinc of Barra ele Navidad (:'-.¡'ickle ancl Collins, unpubl. obs.). Although preferencc for living trccs suggests a high moisture clemand,this has not been subo.;tantiated
through tcsting. Only clealates are knm\'n for th is 'pecies. Should colonie~
containing nymphs he ohtained. we suggest tklt cfforts be macle to
kecp the samplc alivc until :t!ate> clcvclop. Thc'e ,hould be presenccl
in i{)',' ~ alcohol :
:,cncd flight-; abo
ncd: fi dealates, :
de Biología, U;-.,'.-\
:Z: :\fSCollins) VI-~
mary), :! :,olclicrs, 1
same data. l dealat
1 soldiers, 2 nymf
l ~ soldiers, 7 nym
rado Hotel, Barra
(C:oll. l\lSCollins)
comp. A. Emerson
del :\Iaíz . .\[esoph)
Incisitermes platy
similar to emerso1
di ffers in severa! r
:1re "·eakly convex
:¡C); third antenna
(in cmcrsuni it is ~
of the pronotum l
\!Jni . .\landiblc sh:
Vcry littlc is knov
onc in a pine pos
woocl of a hardw,
potentially capabl
t here are no repc
taining alatcs ass<
to our knowleclge
¡, knm1·n from ty
.) soldicrs. Holoty
l-Fl:\0. [.\.\fKH];
I-1~130 . .\fl63. 1
;md comp. ,\. Er
\'lii-:!1-194:!. (Co
11·est bcach El D<
Coll ins) VIII-21-1 1
:\L\.RGI'\"ITER-:\1
unusual -;oldicr e
'íoldier mandiblcs
:rmitidae in the vicinirv
s; C. Anoplotermes; D.
tcterizing this species
termite was first dc>m Guatemala to the
J)' J. Collins (son of
nielad (Collim, unheartwood of stems
line of Barra ele 'N a1 prcferencc for livlt been sub'itantiated
:cies. Shoulcl colonics
efforts be made to
should be presen·ed
1\IcKLF ET CoLLI:'\S.
TERMITES OF
CHAMEL.·\
105
in :{);·~ alcohol along with solcliers and pseudergates. Data on ob-.crved flights also would be useful and important. Spccimens Examinr:d: () dealates, ~R soldiers, 92 nymphs. J\IEXICO: jalisco, Estación
de Biología, Ul\'.\:\I, Charnela. 12 soldiers, 26 nymphs (Coll. D.\Nickle
:Z: :\ISCollins) Vl-2~l-19R1 [N:\I;\'H, CHAi\1]; same data. l dealatc (priman), 2 -.oldiers, JI nymphs [N :\IN H]; same data. 3 soldiers, 28 nympbs
same data. l dealate (primary). ~ soldicrs, 5 nymphs [NMI\'H]; same data.
·1 soldiers, 2 nymph> [Nl\INH]; same data. 1 de al ate (primary queen),
12 soldiers, 7 nymphs. VI-30-l~JR4. [Nl\INHJ; jalisco west beach El Dorado Hotel, Barra de Kavidad. 1 clealate (primary), 2 soldiers, 13 nymphs
(Coll. l\fSCollins) \'I-29-1984 [Nl\fNH]; HOMOTYPE: dealate (det. and
comp. A. Emerson). San Luis Potosí, Cuesta ele los Cedros, 36 km e. Cd.
del :\Iaíz. i\Iesophytic Oak arca. 2 dealatcs (Col!. WLN utting) [AMNH].
Incisitermes platycephalus (Light) 1933. lncisitermes platycephalus is
similar to emersoni in size (lateral length of heael 1.69-2.60 mm) but
di ffers in severa] respects: (1) lateral margins of the head in dorsal view
;1rc weakly comcx, and the posterior margin is distinctly rounded (Fig.
:)C); third antennal articlc is greater in length than articles 4-5 combined
(in em!'rsoni it is slwrter in length than "1-5); anel (3) thc anterior margin
of the pronotum has a eleeper V-shapecl emargination than that of emer\oni. :\Iandiblc shapc ancl clentition cliffer also (scc !'mcrsuni discussion).
\'cry liLtlc is known about this spccics. Light (1933) found two colonies,
onc in a pinc post in a railway cattle guard, the other in the dry sound
wood of a hardwood stump ..\lthough he suggestcd that this spccies is
potcntially capable of hecoming a pcst of structural timber, to date
there are no repon~ of infcstation. As with nigritus and emersoni, obtaining alatcs associated with solelicrs would be an important aeldition
to our knowlcclge of Lhc tcrmite fauna of the Chamela area. This termite
i-, knm\·n from typc material collected in Colima. Spccimens Examinccl:
.) -,oldicrs. Holotype, MEXICO: Colima, ~dadrid, nr. Colima. l soldier.
1-10~\0. l.\:\fi'\H]: P.\R.\TYPE: .\lEXICO: Culima, :\Jadricl, near Colima.
I-l~l~O. l\II6:l. l soldicr (Coii. SFLight). [A:\INHJ; i\fETATYPE (dct.
;md comp . . \. Emcrson): l'cracruz, Potrero Viejo, 10 km. e. C01·doba.
YIII-21-194:!. (Col!. Rl\1Cole). # 22A. I soldicr [.\.\Il\'H]; jali5co, woocls.
\\·cst bcach El Dor:tdo Hotel, Barra de !\'"aviciad. 2 soldiers (Coll. .\fSCollins) VIfl-21-1%8 [:\.\ll\'Hj.
:\IARGI~lTER:\ff.S l\.rishna l%1. This gcnus "·a., erccted for the very
unusual soldicr ct>lc o[ thc monotypic species, hubbardi (Banks). The
soldier mandibles are distinct, and the massively enlargecl third antennal
FoLIA E;-.;Toi>IOL. M rx. 77
lU6
::\'iCKU
( EJ88)
articlc ¡, unlike any
~cnatiw in gcncric e
mes. thc m a in diffcr
aralia.
l \'
f·
\
j '
2 mm
"\larginitcrmcs huhh
'>tinctiYc ami ca-,ily se
The third antennal al
than articles four tlm
in rapid spccies reco¡
bare, with no long s
nac han~ Hi articles.
Thi-, monotypic gcm
,outhern extcnsiom ;
demand and frcc¡ue
tolcrant of high cm
. . ound \\·ood in cxp'
ClelU'i o,kc]etons. c\Ja
rucson, .-\rizona are
uf .\ugmt (:\'utting
~tre<t'i . •\Iargin itermc;
E
-,outhwc-,tcrn Unitec
F.\a/1/ined: 7 soldier
r_;ía. F'\.\:\1, Chamt
IH:\ickle & \ISColl
Ltdo Hotel, Barra d
r:\ \I:\'H].
Fig. :). Dn·\\·ood tctmite ~oldicrs. hcad and pronotmn, dorsal 'ic\\ . . \. B. lnri.1ilcnncs
rwuginipennis. long- aml short-hcadct! fonm; C. l. platyrcpilfl!lll: D. /. cmcrsuni; E, l'.
.\'eolcrmcs joutcli, long- and short-hcaded iorms; C. Cairalilcrllln poiTÚW/11\. (l'igs.
:,\. ll, C. ll and G modifictl frorn Light 1933).
CALCARITERl\IES
nnitl'niii'S are casi!;
phragmotic: C~) fror
ami (3) anterior tit
'pun. lmagoes are d
mm¡ compared witl
nymph mamlible-; al
]c,, pointcd and so
plilied. \\·ith the :\J
'Fig. 3D). Spccies o
of Jncisitcnncs, Cry
fur t he arca.
l
XIcKLE ET CoLLI:\S. TER:\llTEs oF CHAMELA
107
:trticle is unlike any other dry-wood tcrmite. The imago is more con.,ervativc in gcneric characters and closely resembles imagocs of Incisiterme.l, the main clifference being that .Uarginitermes imagoes lack tarsal
arolia.
u
-
'
u
~
, '
'
~
~
Yi<"W . . \, fi. ]nri.lilrnncs
us; D. l. emcrsoni; E. l.
ermes pan•itwtw. (Figs.
\larginitenncs huhhanli ( Ban ks) 1~I:ZO. Soldiers of th is species are distinctin· ami ea~ily separated from other species of the rcgion (Fig. 7C, E).
Thc thinl antennal anide is heavily sderotized ami greatly enlarged, longu
than articles four through nine combined. :\fandible dentition is also u-;dul
in rapid '>pecies recognition (Fig. GG). The ;date ¡, light brown in color.
bare, with no long 'etae on the head or pronotum (Fig. 9C, D). :\ntennae have Hi ;trticJe,. \Ving' (Fig. 1C) are similar to those of Incisitcnnf's.
Thi-; monotypic gcum is distributed throughout the Sonoran Descrt \1ith
,outhern extemiom along the we-;t coast of :\Jexico. lt has a low moislllre
demand and frequently clamages 'tructural timher. In addition. it ¡,
tolerant o[ high cnvironmental temperaturC'; ancl may be found in dry
'ound woocl in expmed arcas. \Ve sampled severa! colonie-; in sagu<tlü
cactm ,keletons. /\lates are nocturnal in dispersa! flights, which in thc
rucson, .\riwna arca are known to occur from :\Jay through the middle
of .-\ugmt (:'\utting 1%~J). Colonies tcntl to be numerous in favorable
;u·ea-;. Jiarginitcrmes huúúardi is common in thc Sonoran Desert from
'outlm-c~tcrn Unitecl States into .\Iexico a-; far as Colima. Spr'cinu·n1
Fxrunincd: 7 'oldiers, ll nymphs . .\IEXICO: jalisco, Estación de Biolo~ía, U.'\.-\:\J, Chamela. In cactus 'keleton. 1 soltlier 11 nymplh (C:oll.
D.\Xickle & .\I'>Collim) \'I-:Z~J-19íH [i'\'\JI\'H]; 1mods, wcst heach El l>or;tdo Hotel, Barra de :\'avidatl. 6 soldiers (Col!. :\ISCollius) VIII-:23-1%8
['\:\1:\H].
L\.LCARITERl\IES Snydcr 1925. Soldier' of species of the genus Calcarikmii'S are ea~ily rccognizable on the basis of 3 characters: ( 1) hcad
phragmotic; (2) frontal margin of head deeply clcft as seen in dorsal view;
ami (3) anterior tibia with one greatly cnlargetl and two smaller apical
\pun. Imagoes are dark Lrown 1rith sm;dl compound cyes (diamctcr ca. (J.~)
mm) compared with head length. .\ntennae ha ve 13- H artide>. Imago j
nymph mandibles are distinctive, with marginal teeth of the left mandihlc
le., pointed ami somewhat fused (Fig. :ZD), and wing venation is simpliiied. \\"ith the :\[ vcin sclerotiletl ami running very close to the R'i
¡Fig. 3D). Specie-; of this genus are found in damper wood than species
of lncisitcrmn, CrytjJtotcrmes, or ,\Iarginitcnncs. Two species are Iisted
lor thc arca.
IUS
FoLIA E:-iTmroL. l\IEx. 77
A
?\ICK
(1988)
(1918), was reportE
have not seen his Sf
lis, not Calcaritennt
in Chamela. Howe'
of CryjJtotermes fat
spccies have a qua·
rectcd to the enlar
been overlooked. 1
Bcsidcs Bank's typ
collected only onu
8
\
D
)\
~
\
G
H
CRYPTOTERM~
1
J
r
l i". li. ~Iandi!Jic dentition of tcrmite soldiers. A. ;-.;,·oternii'S joillcli: 1\. Incisilerme;
marginif!e1111is (long-hcadcd form); C. l. marginipennis (shon-headetl form); D. l.
11ir:1itus: E. l. ,blatyrejilwltts; F. l. emersoni; G. !viarginitenncs hubf,ardi.: H. Amitennes
en_,¡¡,·,; I. .4. CJ)'j!todon; J. Iioj>lotermes amJ!l1ls.
Cdctritermes paninotus ( Light) J9j~l. Soldier> of C. jJaruinotus ha ve
n·< t:tgulatc, elongatccl hcacl whiclt are l.llx longcr than wicle (Fig. :JG).
Vcmbch of thi> specie-. are not commonly cncounterecl, but shoulcl
he sougltt in damp. but -,ouncl. wood. Recorcls of ~date' woulcl he usdul.
Calcariter!l;cs parvinotus is knmvn only from type material collectecl in
Coiima.
Calcuitermcs longicollis (Ban k<,) 1~¡¡ 8. Soldiers of C longicollis ha,·e
quaclrate heacls, wíth a lateral length of the heacl subequal in climension
"·ith thc wiclth. This species. first reported from Panama bv Banks
tive, phragmotic h1
ri tcnnes in the foil
cntire, not deeply t
lar in size. The im<
the dcntition of th
This tropicopolitaJ
termite) capable o
lcaks a'i well as spe1
high hut intermitt
pcsts of structura
-,oldicrs are uscd 1
connccting passag
samples of these tt
sound, sometimes
uninjurccl soldiers
Cryptotermes brev
rugose and convo
hchincl thc fronta
in lcngth, with lE
a 'mrld-wide dist1
ancl their content
known. Its preser
frorn the accumul
l v cxt rnclecl from
pcllct-; is scalcd l
visible on lighter
pcllcts should ind.
ac.cmpted, and tl
l)
·s jollleli; 1\. lllfisilermes
tort-headed form); D. J.
hubhardi; H . .4 mitennes
C. fJaruinotus haye
than wide (Fig. :JG).
mnterecl, but should
ates woulcl he useful.
material collectecl in
· C. longicollis ha\C
1bequal in dimension
1 Panama by Banks
.i\ICKLL ET CoLLI!\s. TERMITEs oF
CnAMELA
109
(1918), was reported from Western :l\Iexico by Light (1933). Since we
have not seen bis specimens (which he described as Cryptotermes longicollis, not Calcaritcrmes), we are unable to wrify the occurrence of this species
in Charnela. However, it is possible he could have misidentifiecl soldiers
of Cryptotenncs fatulus calling them Cryptotcnnes longicollis insteacl. Both
species haw a quadrate, phragmotic heacl, ancl unless attention were directed to the enlarged apical spur on the foretibia, it might easily have
been overlooked. The alate for C. longicollis has not yet been described.
Besicles Bank's type material collected in Panama, this species has been
collected only once, in Sinaloa by Light (1933).
CRYPTOTERMES Bauks 1906. Solcliers of Cryptotermes also llave distinctive, phragmotic heads, but cliffer fram the phragmotic soldiers of Calcaritenncs in the fallowing respects: (1) the frontal riclge of the forehead is
entire, not cleeply emarginatecl; ancl (2) all tibia! spurs of forelegs are similar in size. The imagoes are easily recognibecl as belonging to this genus by
the clentition of the manclibles (fig. 2E) and by 'ring venation (Fig. ~3E).
This tropicopolitan genus contains species (e.g., C. brevis, the furniture
tennite) capable of living in woocl completely protectecl from rainfall or
Ieaks a-, well as species (e.g., cavifmns) found in sound dry woocl in are::ts of
high but intermittent rainfall. Seyeral members of the genus are major
pests of structural timber. The phragmotic stopper-Iike heacls af the
~oldiers are used to prevent movement of attackers through the narrm''
connecting passageways between inhahited chamhers. Collecting gaod
samples of these termites is often a teclious process, since colanies accur in
sound, sometimes wry hard wood, and much care is required in securing
uninjurecl solcliers. Two species accur in .Jalisco S tate.
Cryptotermes brevis (Walker) 1853. The solclier of this species has a dccply
rugase ancl canvoluted, highly phragmotic !Jcad aml a deep depression
hehincl the frontal riclge (Fig. 8A, B). \Vinged adults are about 10 mm
in Iength, with 16-18 antennal articles. This is a majar pest species with
a "·orld-wide distribution. To date it has been found only in buildings
and their contents; its accurrence in natural environments is as yet unknown. Its presence in lmildings ancl their contents is usually cletected
from the accumulation of small piles of fecal pellets which are periodically cxtrucled from inhahited gallcries. The hole created ta extrucle the
pc!Iets is scaled by a smooth film of salín and ''-'Oocl residucs and is
\'Ísible cm lir~hter shacles of wood. Presence of these openings or piles of
pclleb slwuld inclicatc the need for a thorough treatment befare rcpairs are
at<:cmptcd, ami the application of soft ph woacls should be ayoicled.
!lO
FoLIA
E:-;To:.\101..
l\1 EX. 77 ( l98H)
Cryptotennes fatulus (Light) 1933. The on1y naturally occurring species
of this genus in the Charnela region is C. fatulus. Soldiers have a finely
crenelated raised frontal ridge and subquadrate head shape (Fig. 7E, l'}
Lateral length of the head ranges from 1.1 to 1.2 mm. A.lates of fatu/u;
(Fig~. 9A, B, G) are a much deeper brown than the larger alates of
brc;ús. Both have light brown, iridescent wings with similar wnation
(Fig-. 3E). Head length of fatulus ;dates is 1.13 mm (cf. brcvis, 1.3 mm)
al1<1 total body length is 6.5 mm (cf. brcvis, 10-11 mm). Light (1933) collect
cd ;dates of C. fatulus on l\Iaria Madre Island on l\Iay l!í. This specie,
occurs in dry sound wood, such as downcd timbcr, fence posts, and outbuilding foundations. Thc nest galleries comiH of 1arge chambers coBnectcd by narrow, round tunncls. Additional data in dispersa! flight times
would be a useful addition to the currcnt body of infonnation on the
termites of thc Chamcla area. SpeciJnens Exmnined: 5 imagocs, 8 ~oldiers.
·lH nymphs. :\IEXICO: jalisco) Estación de Biología, Ui\'Ai\1, Chamela.
1 ala te, ~ de al ates (primary?), 3 soldicrs, 21 nymphs colony Gl (Col!.
DA:\'ickle & l\ISCollins) Vl-~9-198'1 [1\':\INH]; same data. l alatc. 3 soldier-..
21 nymphs [Nl\11\"H]: 'iame elata. 1 clealate (primary?), 2 soldiers, 6 nymph-..
\ I-:W-I~l81. [N:\f;\'HJ.
:\!CKI
on the clorsoanteri(
has mandibular deJ
3G. The imago he
which oftcn conceal
light brown. Heter,
or cxposed wood
wooden structures a
for a closely related
arca, occur from Jt
a ureus aureus is fo1
tbe species range, eJ
occupics lowland s
Venezuela. Specime
fisco, Barra de :\avi
(Coll. DA:\ ickle) \'.
[N:\Il'\'H]; same dat
mela, UNAi\I 12 S(
29-l~)R} [X\INH]; S
tíO nymph~ [NMNI
dicrs, 2:) nymphs [
\ '1-30-1981. 2 soldic
RHt:-IOTFR"JTIDAJ·: Subtcrranean Tcrmites
HETEROTERMES Froggatt l98ti. Soldiers of Hcteroten11c:, lta\e yello\1·
clongatecl, rectangulatc heacls with vcry sma!J fontanelles, situated betwecn
nro ridge-> on the clorsoanterior region of the head capsule. Mamlibles are
elongatccl, slcncler, and apically recurved and pointcd. Inner margins of
the mandibles lack marginal tccth, although ba'ia!ly thc margins may be
wcakly serrated. Imagoes are light brown or yellowish, with elongated hcads
and small circular fontanclles. :\Ianclible dentition, (Fig. 2F) and \\'ing vcn:ttion (Fig·. ~K;.) are uscful in gencric rcc:ognition of thc imagocs. Specic,
of this gcnus usually are founcl a-; Ltrge colonics of small tcrmites
liYing in wood, with suhtcrrancan nest'i frequently conncctccl to fceding
,itc' by co\crcd runways. \Iany species fcccl on structural timber, and
,mne are comiclcrecl to be major pesb. Only onc spccics oc:ctu'i in thc
arca surrounding Chamela, but locally it is oftcn a maJor pe't.
COPTOTERMES
tinctive oval or p)
tanclles projecting ;
elongatccl, and apic
are untoothed. Ima
articles. l\Iandibles
useful in recognizÍl
in the field from ti
of thick liquid at t
i-. easy in both fiel
'iclcr:thlc more cliff
prcsentcd morphm
'olclicr ctste, but
idcntification of sir
is Ji-,ted for the an
Heterotcnncs aureus convcxinotatus Snydcr l9:Z1. Soldiers o[ this speoe:i
kt\'C goldcn to ycllowish or ambcr-colored, rectangulate hcads. :\fandiblc' ;trc 1.2x longcr than hcad 11·idth and lack tecth along thc inner
margms. Thc fontanclle is cviclent as a small pinhole betwecn two ridges
Coptotermcs crassu
Jargc antcriorly-clil
( Fig. 1OA. B) Fort1
S)
rally occurring species
Soldiers havc a finely
id shape (Fig. 7E, F).
mm. Alates of fatulu.,
the larger alates of
with similar \·enation
1 (cf. brevis, !.3 mm)
1). Light (1933) collect
May !:J. This specie,
fence posts, and out. large chambers con1 dispersa! flight times
f information on the
5 imagoes, R soldicr,,
ía, UNA!\!, Chamela.
)hs colony C-:1 (Col!.
ata. l alatc. 3 soldicr-,,
, 2 soldiers. 6 nymph-;.
::\ICKLF
ET
Cou.I:-;s.
TEK\riTEs
oF CHA\IELA
111
on the dorsoanterior portion of the head (Fig. IOC, D). The imago
has mandibular dentition as in Fig. 2F, ancl wing venation as in Fig.
3G. The imago head is elongated, clothed with numcrous long setae
which often conccal the fontanclle (Fig. 1 lA, B): body color is golden to
light brown. Hetcrotennes aureus cunvexinutatus is common in buried
or exposed wood ami often causes considerable damagc to untreated
wooden structurcs aml storcd products contained therein. Dispersa! flight'i
for a closely related subspecies, H. aureus aureus in thc Tucson, Arizona
arca, occur from .J une through Septembcr (r\ utting 1~Jii9). Heterutcnnrs
aureus aureus is found at higher elevations and in northern portions of
the spccies rangc, extending into "\rizona, whilc 1-f. auu:us ronvexinotatu'
occupics lowland sites and cxtends further south to Panama and into
Venezuela. Specimens E.\aminerl: 62 soldiers. 100 nymph'. :\JEXIC:O: jalisco, Barra de l\'avidad, west bcach El Dorado Hotel. 2 soldie!";, S workcr:(Coll. DA:'\"icklc) \'f-29-191\·l [N~INH]: same data, 20 soldicrs. 200 nymphs
[N;\Il\H]: same data, 3 soldicrs, 100 nymphs [N~INH]; Estación de CIJamela, UNAl\I 12 soldiers, 9 nymphs (Coii. DANickle & .\ISCollins) VI29-1981 rNMNH]: same data, 12 soldiers, 11 nymphs [N;'\INH]: 2 solclier-,.
60 nymphs [Nl\INHJ; same data, 3 soldicrs, ·1 nymphs [¡\':\fNHj: 5 ~ol­
dicrs, 25 nymphs [N.\INHJ; 2 soldicrs, :'í nymphs [NMI\H]: samc data,
\'I-30-191\1. 2 soldicrs, 6 nymphs [N:\1:\"H].
mi tes
rotennc:, ha \'C yello\1·
!lles, situated betwccn
1psule. Mandibles are
ed. Inner margins of
the margins may be
with elongated heads
Fig. 2F) and wing YC. the imagoes. Spccie,
!S of small tcnnitcs
connected to feeding
ructural timbcr, aml
;pecies occurs in thc
major pest.
,Jdiers of this species
tgulate hcads. i\Ian!eth along the inncr
~ between two ridge>
COPTOTERMES \\'asmann 18%. Soldiers of CojJtotermes have dj,.
tinctive oval or pyriform heads, with enlarged oval to triangulate fontanclles projccting anteriorly just abovc the clypeus. ;\Iandibles are slendcr.
elongatcd, and apically rccuned. anc1 the inner margins of the mandiblc,
are untoothed. Imagoe-> have broad subcircubr hcads with !R-21 antennal
articlcs. Mandibles (Fig. 2G) and wing venation (Fig. ;',F) also are
useful in recoguizing membcrs of this genus. Soldicrs are easily recogui1cd
in the ficld from their habit when disturbecl of extrwling a milky globulc
of thick lir¡uid al the front cncl of thc head. \Yhcreas gencric recognition
is casy in both field aml laboratory situations, spccies recognition is con;.idcrablc more difficnlt. Light (1927) ami Roonwall and Chotani (1962;
prcsentcd morphometric methods for separating specic-;, IJased on the
,oJdier castc, lmt interspccific o\·erlap of such charactcrs makcs specic'
idcntification of single inclividuab a clifficult task at bcst. Only one specic-;
is Ji,tcd for thc arca, but it is a potcntially important pe't specics.
Coptotermcs crassus Snydcr !922. Soldier hc:Hl shape is pyrifonn, with thc
large antcriorly-dircctccl fontanellc ~ituatcd dirccth' bchind tbe clypcm
(Fig. lOA. B) Fortunatcly, since tliis is thc only species of the gcnus in
ll~
FouA E;o.;To.\IOL. :\lt:x. 77 (1988)
the Charnela region, species recognition is not a problem. Alates are dark
brown to nearly black, with a broadly roundcd hirsute head, as seen in
dorsal aspect (Fig. ll C, D). Fl ight pattern is diurna!; Light (1933) recorded dispersa] flights during July and August. This species shares with
others in the genus the ability to attack both living ancl dead trees
ancl to live either in subterranean nests or in wood free of soil contact.
Colonies may become very large, aml damage to wooden structures is
often severe. Cuptotenncs crassus has been rcported in :\Jexico, Guatemala, Honduras, ancl Panama.
;\ IC
~hould be recorclc(
mela area, althou¡:
------
Carton-Nest Termites, Dcsert Termites,
Soldierless Termites and Others
TER:\!ITIDAE:
FOLl
This i~ the mmt spccialized family of tennites, ancl members of the
group are often thc most obvious elcments of the termitc fauna, from
the habit of many specics in constructing conspicuous carton nests on u·ees,
stumps, or buildings. In adclition, howevcr, the family includes manv
specics which are subterranean and inconspicuous.
Las lámina
(págs. 112 y
tercaladas er
AMlTERMES Sihestri 1901. Solclicrs of Amitcnnr·s have oval to rectangulate heads with largc oval to circular fontanelles, situatecl anteriorly
on the raisccl front of the head and surrounded by numerous long setae.
:\fandibles are usually less than 0.80x as long as the head width; however,
they are clongated, symmetrical, usually strongly recurvcd from base to
tip, and armed on the inner margins with a single tooth which may he
moclified as an elevated cutting bladc arising near and extending to the
apex. Imagocs are usually dark brown to black, and relatively small (rarely exccecling 12 mm). Head shape is usually subcircular, ancl the fontanclle
is conspicuous, rouncled, or elongated. !mago manclibles (Fig. 2L) and
wing venation (Fig. 1E) are useful cliagnostic features. Foraging· groups
of thcse termites are usually found uncler large stones, in and under old,
dricd cow clung. in old dead stumps, other partially-buried wood, and in
dead fallen branclws. Soldiers should be collected rapidly upon exposure,
becamc, whcn disturbed, thcy tend to rctreat quickly into unclcrground
galleric-;. This gmup has been clifficult to charactcrize fully. primarily
bccathC of the clifficulty in collecting soldien, ami abtcs together. If
'vin¡,;nl fonm are observecl during dispersa! flights. efforts should be
dircctecl toward locating the sourcc of the swarm. and somc workcrs ancl
solclicn from thc source ncst collccted with the s11·armers .. \s with othct
s¡wc;'~s. the time of clay, the date. ancl associated wcather conditions
of this species are
tceth arising near t
brmrn, about 1:? 1
large, oyal, nearly
Defcnse in A. ¡¡•he
CISC ,~·it!J aJl Othe
manclibles whose d
t ion from the fror
:me! collects arouncl
dccl, coati11g an att;
after contact with
lenged, ants will nc
.:\JCKLL Er CoLLI:'\s. TER:\llTEs oF CHA.\1ELA
roblem. Alates are dark
lirsute head, as seen in
urna!; Light ( 1933) refhis species shares with
living and dead u·ees
10d free of soil contact.
o wooden structures is
~ted in :\fexico, Guate-
113
should be reconlcd for the fl ights. Fiw species are known from the Chamela area, although species names haYe beeu confirmed for onlv two of
FE DE ERRATA
ert Termites,
ers
FOLIA ENTOMOLOGICA MEXICANA No. 77
;, and members of thc
1e termite fauna, from
us carton nests on trees,
family inclucles manv
Las láminas de las Figs. 7 a 15 que deberían entrar en este lugar,
(págs. ll2 y 11 "9.) , por un error de encuadernación f¡uedaron in-
;.
terca1adas entre las págs. 512 y 513.
mes have oval to rec~lles,
situated anteriorly
'Y numerous long setae.
1e head width; however,
recurved from base to
le tooth which may be
r and extending to the
d relatively small (rareular, and the fontanclle
mdibles (Fig. 2L) and
ttures. Foraging groups
mes, in and under old,
ly-buried wood, and in
rapidly upon exposure,
[ckly into undcrground
.cterize fully. primarily
m el ala tes togcther. If
~hts. efforts should be
and some workcrs and
~-armers. As with othcr
.ed wcathcr conditions
11 JUUl ot V.!J-l.l. mm. The most characteristic features
this ~pecies are the robmt mandiblcs with well dcveloped, peg-like
tceth arisiug near the basal thinl of the inner margins. The imago is dark
brmm, about 1:2 mm in lcngth, with a slit-like fontanelle. Ocelli are
Jarge, m al, nearly half as long as the cliameter of the compound eye.
Defense in A. ¡¡.¡zn:lcri has received some attention. As is probably the
case with all othcr '>pecies of thc genus, solclicrs have well developed
mandibles wJw,e dcfensive function is augmcntcd by a chcmical secretion from the frontal gland of thc hcad. This secretion trickles down
ami collccts arouncl the bri.-,tles of thc labrum. The lahrum can oc extrudccl, coating an attacking insect with the -.ecretion. Ants groom Yigorously
aftcr contact with this material. Furthermore, when experimentally challenged, ants "·i]] not cross a barrier coatcd with solclier secretion, indicat-
... "'"" '"'u ,, ''c.tu
o[
188)
'roblem. Alates are dark
,:~""tp hFad. as seen in
d reJa u ve~ y """'"~ ,
ular, and the fontanelle
mdibles (Fig. 2L) and
ttures. Foraging groups
me;, in and under old,
Jy-buried wood, and in
rapidly upon cxposure,
[ckly into underground
:cterize fully, primarily
md alates together. If
~hts, efforts should be
and some workcrs and
\'armers. As with other
.ed wcather conditions
:'\" JCKLL Er CoLLJ:\s.
TER.\IITES oF
CHA:\1ELA
113
should be reconlcd for the flights. Five species are known from the Chamela arca, although specie~ names have bcen confirmed for only two of
these forms.
Amitermes cryptodon Light 1~130b. This is the most atypical of the species
of Amitcrmc~ of the Chamela arca. Soldier mandibles are elongated,
near!y as long as the head capsule, ami only >reakly rccurved along its
lcngtl!. ] lte tooth ¡, ~ituatecl at the base of the mandible, directly beneath
thc apex of the hbrum. ami extending as a cutting blade to the apex of
the mandible \ F ig. l :)C ). TlJe imago Giste is unknown for this species.
Light ( l930L) clc,aibed thi~ specics from severa! colonies collected at
Jala, 21 km\\'. of Colima. and ~Iaclrid, 28 km \'V. of Colima. \Ve collccted
onc smaJJ ,;¡mpie ;:t Barra de :\avicLtd, on the coa'lt just north of Chamela ..-\11 s;nnpJe, 11cre fouml in decaying >mocl in ancl on the ground. Covercd
earthen rumra;s 11·ere macle on the wood being attackcd. Specimens
F.\amincd: í solclier-,. lO 1\'0rkers. "\IEXICO: on ro u te 15, between Tcpic
and Guadalajara, cm1· chip at '1 000 ft. 6 soldiers, l worker (Col!. l\ISCollim) \'lll-l2-E16R [:\.\f."\H]; Jalisco, Barra de ?\avidad, 1rest hcach El
Dorado Hotel. l solclier, 9 workers (Coll. DANickle) [T\l\I:!\'H].
Amitcrmcs cnsifcr Light 1930L. Light (1933) reports this specics m Jala,
near Colima 1Colima State). \\'e haw not collectcd it in Jalisco. The soldicr
h:ts long, weakly-cuncd mandibles with cutting shelves locatcd along the
distal thircl (Fig. 6G). The alate is unkno\\'!1 for this species.
Amitermes whccleri (De,neux) 1906. Amitcnue.1 wlu:elcri has the largest
soldier of the Amitcrmes specics o[ the region, with a head length of 1.21.1 mm :md a heacl 11·idth of 0.9-l.l. mm. The most characteristic features
of this species are the robmt mandiblcs with well dcveloped, peg-like
teeth arising near the ba-.al thircl of thc inner margius. The imago is dark
brmm, about 1:2 mm in length, with a slit-like fontanclle. Ocelli are
Jarge, onl, nearly half as long as the cliameter of thc compound cye.
Defense in A. <chcclcri has received some attention. As is probably the
case ~~·ith all othcr -,pecies of thc genus, soldiers have well developed
manclibles whose clefensiye function ¡, augmented by a chcmical secretion from the frontal gland of the heacl. This sccretion tricklcs clown
and collects arouncl the bristles of thc labrum. The lahrum can be extruded, coating an attacking insect with the 'ecretion. Ants groom Yigorously
after mntact 1rith th is material. Fmthermore, whcn experimental! y challenged, ants will not cross a barrier coatecl with soldier secretion, indicat-
111
FoLL\ Exro.\IOI. ,\lis. í7
:\IC
(l~lK0)
ing that the suhstance i~ repellent in addition lo irriL1ting, and acL'> as
a deterrent to predation (Scheffralm el a l. 1~!K~l )· .\nalpi-, of the solclier
secretion in cliffercnt populations of A. 7l'hr·cleri in Cdifomia rcyealed
that the :,ecrctiom of :,oldiers of different populatiOih could he highly
variable, with thc cxception of a 'ingle component, helminthogermacrcne, an isomer of germacrcne, \\·hich "''" found in thc -,ecretiom of ~di
A. ¡¡•!lec/ni soldiers stuclicd (Scheffrahn 1'1 o/. ]q01i). Thi> consistencY
suggests futurc utility as a chemotaxonomic char:H ter. Soldier> are comparatiYcly rare, comprising ~~ smaller proportion of thc toLII colony t han
in uuny other genera. Schcffralm ct al. (ElK:l) -,uggc-,t that eycn though
fe"· in numher, the -;olclicr;, could dcfend a colony c.::ffectiYcly. if the
tcrmitcs foragecl undcrground ~tml comtructcd solicl nc'>h with tough runW~I'"'· Thc soldier-, coulcl function importantly in hlocling nc'>l entrances. \Vec;,ncr (l~Hi:>) :,tate-; that flighh o[ thi, ;,pecic-, in .\rizona ou:ur
;¡t du:,k following c~1rly ;,uJllllllT >hmrcrs. Thi-, ¡, thc mo\1 wiclc,pread
spcl ÍC'> ul Alllilr'rltll'\ in :\'ortl1 .\mnica, occurring from Calilurni~1 tu
Tcx;¡-, in the llnitcd States, :,outh to Colima and J;t!i-,co StalC'> in \lex.ico.
sp~·cimcm EYa111incd: ;;oo ~oldicr'>. :"¡()() \wrkcr,. \!EXICO: Jalisco. Estaciún de 1\iologí:I, LT:\'.\\f, ChamcLt. 7 '><Illlplc': ;)()() '>olclic:r'. :"¡()() \wrku-..
in tot:t!. (Col!. IL\.'\icklc "-: \ISCollins) \'1-~CJ-1~101 [:'\\1:\H].
.\mitcrmes spccie-, 1 (ma; be parvulus Light EU~). lt ,,;¡-, not po·:siiJlc
to compare thi:, spccie-; collccted al Chamcla \l·ith Light\ matcri;d ot thc
C;difornia -,pctÍc;, pan 11!ius, IJut thc soldicr-, of our material agrcc doseh
wit h Light ·, ( 1(1:\~) dc-,cript ion for parvulus. I.cngt h of head capsulc i'
;dJOut l.O mm, ami mamlihlc shapc :md dcntítion (Fig·. 1:1.\. n) agrcc
do-,cly jJIII"i'llills. If t\li-; i-, A. j)(lnmlw. our rccod-, hcrc report a m;Ijor
r:mgc cxtcn-.ion for thc spccic'. -,iuce j)(ln'ulz!l ¡, currently knm1·n onh·
from C:d iforni;I.
.-\mitennes 'Jll'l ic-, ~- Thc ,o!dicr of thi, -,pene-, J¡;¡-, the !olio\\ ing c]j,_
tinctin~ ch;IrarlCJ'< (l) dypcm hilohcd: (:2) nurgin:d tooth sm;t!l, pcglike, ;¡ri-,ing at b;~;,~¡\ thircl o! mamliblc. (?)) a ,,·cakl; dcn:lopcd cutting
edgc arising at marginal tooth ami cxtending to :1pex. (Fig. l3E, F).
\!ore material ¡, nccdccl tu confirm thc idcntity of thi' 'iwcics.
G!..;ATH.\MITER:\IES Ligllt 19~\~ . ."oldier-; ol
(;¡¡alluunitr·rn¡cs are SImilar to those of Alllifl'lllll'.\ cxccpt that thc m:III<liblc-; ~1rc grcatcr than
O.Kx width of head. typically ;dJUut l.Ox widtil. ;md nearl) straight
t hrouglwut t:Jeir lcn:;th. Thc imago ¡, ];u gcr than f01 Ami t !T/111'.\J greatcr
than l~ mm, in mmt C\SC'>. \\'ing Ycnation (Fig . .fB) ;lllcl imago mandiblc
dentition (Fig. ~ i
t hose of A 111 itcnnt
h o m t he arca.
Gnathamitermcs t1
casilv recogni1ed b
11.-ig. 1·1D). The dil
'Jll'( ic-; has rc'>nltec
dn 1~~ I(J; .\raujo 1
í n t he Chamcla ar
Ltkc this opportun
hot h e astes '>O that
of this genus. TI
'ullcctcd by us in
Gnathamitcrmes p
cugni1cd by the si
1 lE). Light (19!l3
i t ;¡-, t he '>pecie-; i
íl/1'\ /1.1 ).
Gnathamitennes s¡
of t!Ii-, fonu. lt m
'll( ]¡ a judgement,
frmn t!Ic rcgion. Ir ion are distinctÍY<
f !OPLOTERJ\IES
r 1te -.o!dicr ancl w
lJCJth in \lcxicu ar
[ lo¡¡:otermes amp:
c;¡-,il) distinguisllec
brgc (~.:~ mm in
¡'' c.,-,cd ;¡¡ea in th
unclle (Fig. !~E,
¡, 1 uhmt "·ith a si
unknown for this
:tnd usually llave (
taincd in the vol
a~ the "brge. blac
log and largc tim
1\:JcKu
rritating. and ach as
,nalpi'> of tbe soldier
n California re\·ealed
iom could be highly
ent, hclminthogerma1 the secretiolb of all
186). Thi~ consistencY
ter. Solclier:, are comthe total colonv than
sest that cven though
,ny effectivcly.. if the
nesb 11·ith tough runin blocki ng· nc-,t cnecics in .\rizo11a occur
the mo~t widespread
g from Calilorni~t to
lisco Statcs in :\fcxico.
EXICO: ]aliscu, Estasoldier:-. :iOO \mrker-..
ET
CoLI.l:\~. TLR\lln~~ or- CIL\\lt.LA
115
de11tition (Fig. ~ "\!) also scnc to distingui,f¡ membcr'> of this gcnu'i from
thosc of Alllitcmlr's. \Ve havc idcntificd thrce '>pccics o[ (;nat!w111itennes
from t he are a.
Gnathamitermcs tubifonnans ( Buckley) l H(j:J. Soldiers of thi, specie~ are
c~t,ily rccognized by the cutting shclf 011 thc apical third of cach mandihle
(Fig·. 14D). Thc difficulty in associating· wingcd fonns with soldicrs of thi'
'fJCC ics ha-; re'>ttltcd in numcrous misidentifications in thc litcrature (Snydn Pl El; .\raujo 1~l77). Si11cc \\'C ha\'c mlt collcnccl -;oldicrs with imagoc-.
in t he Chamcla ;trca, \\'C choosc 110t to a del to thc confusion. Jn-;tead. we
takc this opportunity to suggest that more collections he m~tclc to collect
hoth ca'>te.'> '>O that futurc work will not add to the taxonomic problcnh
of this genus. This specie'>, k11own previou-.1; only from Tcxa,, was
( ullectccl by us in Jalisco.
Gnathamitcrmes perplexus (Banks) 19~0. Solclicrs of thi-; species are tccognilcd by the shape of thc hcad ami dentition of thc manclihlcs (Fig.
!lE). Light (E!.)3) reponed its occurence in thc Chamcb area, listing
it as thc '>pecie'> infwnatus (now considcred a junior '>Yll011)lll of fJr'r·
1¡J¡·_y /{,\ ).
[:'\ \1:'\H].
). It ,,·a-, Hot pm,ible
Light\ material of the
material agree clo:-cly
¡th of head capsulc i'
1 (Fig. !3,\. U) agrec
; hcrc report ;¡ m;tjor
cmrently knm1·n only
has thc lollm1·ing dj,_
;in;d tooth sm;dl, pegakh· dewlopcd cutt ing
) z:¡)ex (Fio.
13E, Fl' .
. o
>f this :.pccics.
;nalhamitcrrnrs are ~i­
diblcs are greater th~m
1, ami nearly stra ight
for A mitcnncs, grcater
B) and imago mandiblc
Gnathamitenncs species l. \Ve have beeu unable lo determine thc '>pecie,
of thi-; form. lt may he new, lmt until \IT haYc more materi;tl to make
sw h ;¡ judgcment, wc '>imply list it as <lllotlwr Cnatluunitcnncs '>pccic'
from the rcgion. Hcacl shapc, bilobcd clype;d mturc. aml mandiblc dcnl ition are di'otincti\'c for thi-; '>fX'cie-; (Fig. JIF).
IIOPLOTER:\IES Light l~l;\.'\. This lnonot,pic genu' i-, knmm onlv from
the -.olclier ami workcr ca,.,tc-;. lt has bccn <ollccted only a fcw times.
both in :\lexico ;me! GuatemaLt.
!lo¡)iotermcs amplus Light l<J:U. Soldiers ol thi.-, unmual tcrmite are
c~hih di'otinguishcd from other tcrmite-; of tlte rcgion. Thc head capsulc i'
Lngc (~_:, mm in lcngth). rectangubtc. ;md honey colored, with a deJ'le,,ed ~trca in the front of thc ltcad m~trking thc position of the fonLlltcllc (Fig. !~E. F). The apcx of thc Ltbrum i-; notchccl. Eacl1 m;mdibk
¡, robu'ot with a single shallow tooth on thc inner margin. Thc alate is
unknown for thi-. monotypic species. The workers are rclativcly large
;tnd usu;t!ly lJayc dark hrown to black abdomens from the material contained in thc voluminous gut, ctusing Light (19.).)) to refer to thcm
:t'i the "Ltrge. hlark-bellied tcrmite." He found thcm fccding 011 an old
log ami large timhcr on the ground: l'\utting (pers. commun.) found
ll6
FoLIA El\TO\IOL. ~h:x.
77 ( l98R)
N re K
thcm under stoncs besidc corn ficlds in Ajicjic; and we (Nickle and
Collins) found thcm to be common under large stones and desiccatcd
cmr dung in pasturcs in northcrn Guatemala. Foraging groups of this
little-knmrn tcrmite can be locatecl by turning oycr large :,tones, especial!¡
in pastures at higher eleYation' than slwreline habitats typical of the
Chamela arca. As with Amitennes collecting, the trowel/tray tcchniquc
works "Til hcre. U sualh' onlv one or a few soldiers can be found with
the foraging workers.
J
'
AXOPLOTERMES ~lüller 1873. The:,e high1y-specialized mcmbers of thc
Termitidac llave clispemecl with thc soldicr castc altogcther. Insteacl, thc
cldensivc functiom are carriccl out by 1vorkcrs, which expel salivary sc-cretion or liquicl gut contents, using vigorous muscular contractions o[ thc
abdmncJL Rclatecl genera in othcr p;trts of thc tropic'i dcfencl thcm-,cl\c,
11'ith clroplcts of fcccs applied to thc hcacls of attackers, or cxplocle tbc abdomcm in thc arca of the attackcrs, releasing Iarge droplcts of intestinal
m!ltcnb. This set of strategics may be quite cffectivc: for imtancc, no :\eotropical .\rmy Ant attacks solclierless tennitcs (Collim, pcrs, obs.). Thc
workcrs of .,OJllC soldicrlcss tcrmitcs are quite aggressive, attacking ants \l'itll
thcir mandiblcs befare applying cligestiYc rcsiclues. This gcnm is in
nced of revi;,ion for Kc11· 'Vorlcl rcpre'ientatiyes. Iclentifications must be
bascd on thc imagoes until 1wrker characteristics suitable for taxonomic
differentiation are available. 'Ving vcnation ( Fig. ·1C) and mandible clentition (Fig. ~H) are the most reliable charactcrs for gcneric iclentification.
.\noplotermes fumosus (Hagen) lR60. Both workers ami alates of thi,
species are readil)' recognizcd b)' mandible clentition (Fig. 2H). A.late'i
haYe Iarge occlli and compound eycs, and thc ocelli are <;eparated from
thc compound cycs by a clistancc equal to the occllar cliametcr. The
fontancllc is inconspicuous . .'\late mandiblcs are e1ongated ancl protru,ive
heyond the apex of thc labrum. Antcnnae are 14-scgmented, and tibia!
~purs are 3:~:2. "'orkers of this species llave a dusty appcarance, 1\'ith
Iight cross striatiom on thc abdomen. At Iower elcvations, these termites
may be founcl most commonly in ami under clead wood. Related spccie-;
at highcr elevations are found in cow chips . .'\lates are founcl only sea,onally; if they are present, a good sample along with workers of the varioLb
sizes prcsent shoulcl be preserved. :\Iuch remains to be learned about tlli-;
intcresting group of tcnnites. A kcy using charactcrs from scyeral region~
of the cligestivc tract has been published for membcrs of this group in
Sout h .\merica (J ohnson 1979).
:\1 ICROCEROTER
manclibles are mod<
tinct rough serrati<
on the so1dier head
haw elongatcd hc2
:\ntennae are 15-se¡
thi> genus are knm1
::\Iicrocerotermes gr;
JI icrocerotcrmcs gr.
head and clongatec
tr·ntrionalis has a 1
mancliblcs (Figs 12
í 1~~~\!\) founcl scptcn
i ng ,¡tes bu t found
f¡ ionalis is abunda
!:racilis is more Ji
llave not bcen descr
havc been obsened
merom, patient cffc
TERMES Linneaus
reco~nizecl in the l
ll'ith a short anter:
;mtcnnae (Fig. 11B
the shapc of the m
the head capsule, v\
<ip:ttulatc (Fig. 14A
diblcs are black. TI
Yicw, large circular
1 I E, F). The fontan
dc11tition (Fig. 2K)
lD ¡ anc! wing colm
;pur~ are 3:2:2, but
'ctae. giying thc ap¡
t\ pes of tcrmites wl
:"ymmetrically twis
ancl proclucing an ¡
'trikc intruders with
c:ollins ami Prcstwi'
ing comhat cfficien<
8)
and we (N ickle a nd
stones and desiccated
)raging groups of this
largc stones, cspecialh
tabitats typical of the
troweljtray techniquc
~rs can be found "ith
:ializecl mcmbers o[ thc
1ltogether. Insteacl, the
eh expel salivary secretar contractions of the
'pies clefend them-,ch e'
~ers, or explode the abe clrop!ets of intc~tinal
~: for imtance, no Xco)llins, pers, obs.). The
ive, attacking ants \l·ith
tes. This gcnns is in
dentifications mmL he
suitablc for taxonomic
iC) ancl mandible den. generic iclentification.
ers and ala tes of th i.,
tion (Fig. 2H). A.late-;
~lli are separated from
ocellar diameter. The
ongatecl ancl protnhive
l-segmentecl, and tibia!
lusty appcarance, \l·ith
~vations, thesc ter mi tes
wood. Related speric-;
are found only seasonworkers of the variott>
, be learned about tili-;
rs from severa! regiom
abers of this group in
~ ICKLE ET CoLu:-;s. TER:\IITEs oF CHA;\1ELA
117
~IICROCEROTERMES Silvestri 1901. In soldiers of Microcerotennes the
mandibles are moderately elongated, flattcned, with characteristically distinct rough serrations along the entire inner margins. The fontanelle
on the soldicr head is small and usually very difficult to locate. Imagoes
haYe clongated heacls with small- to meclium-sizecl circular fontanelles.
Antennae are 15-segmcntecl, ancl tibia! spurs are 3: 1:2. Two species of
this genus are known from thc Charnela arca.
:\Iicrocerotermcs gracilis Light 1933 and M. septentrionalis Light 1933.
Jiicroccmtcrmcs gracilis is the smaller of the two specics, with an oval
head and elongated, finely-serratcd mandibles (Fig. 12A, B), while scptnitrionalis has a rcctangulate head and short, robust, coarsely-serrated
m;mdibles (Figs 12C, D). Nests of both spccies are subterranean; Light
1l ~l~\3) founcl scptcntrionalis building extensive covered runways to feedíng sites but founcl no such rtlll'xay-; for gracilis. iHicrocr:rotenncs septentriunalis is abundant at highcr altitudes (El Rindm, Colima), while
is more likelv• to be encountercd at lower altitudes. Ala te-;
0i' racilis
h;tYC nol bcen describccl for either spccies. Severa! distinct types of workers
haYe heen ohscrved in mature colonics. Since soldiers are not very numerom. patient effort may be rcquirecl to secure :m adequate sample.
TERMES Linneaus 1758. Solcliers of Termes are distinctive ancl easily
recognized in the field. The heacl is rectaugulate, posteriorly rounded,
11·itlt a short anteriorly-directed conical projection located between the
antennac (Fig. 1m, C). The most distinctive soldier feature, however, is
the shape of the manclibles. l\fanclibles are elongate, usually longer than
the head capsule, weakly asymmetrical. clm\·ncurvecl, ancl apically weakh·
~patulate (Fig. l4A). Thc head capsule is usually white, while the mandibles are black. The imago has a broad, rounded, hcacl as seen in dorsal
Yiew, 1arge circular occlli but small-to medium-sized compouncl eves (Fig.
11 E, F). The fontanelle is slit-like, sometimes clifficult to locatc. ~Lmdible
dc11tition (Fig. 2K) is useful in generic recognition. \Ving venation (Fig.
} D) ancl wing color (usual! y dar k smoky grey) are al so helpful. Tibia!
>purs are 3:2:2, but the hindtibiae have one or two spine-like prc-apical
'etac, giving the appearance of 3:2:3 or 3:2:c1. This genus is onc of severa!
t vpes of termitcs whose solcliers ha ve jaws variously prolongecl, frcquentlv
:t-,ymmetrically twistecl, functioning in dcfensc by powerful uucrossings,
ancl proclucing an audible snapping souncl. The uncrossings function to
strike intruders with a powerful force lo knock them away from the solclier.
C:ollins aml Prestwich (El8:l) proYide proccclures ancl guidclines for studying combat efficienc:y comparisons.
:\¡
lit\
FouA E'>TO\IOI ..
\lix. 77 (l~JR8)
Termes panamaensis (Snydcr) 19~3. Colonies ol this species are usual! y
lound in dead, partly huried st umps. cspccially coconut palms. Obscrvations of dcfcnsive bchavior would be of spccial intcrcst, covering such
points as perccntage of soldiers in reprcsentative colonies; thc teclmiquc,
usccl by soldiers in prcventing cntrancc of woulcl-be prcdators into clamaged ncst arcas, ami the most common predators. Vouchcr specimcns of
both the tennitcs ami the attackers should be prescrvcd. This species
has bcen collectcd in \fexico, Guatemala, Costa Rica, ami Panama. Spccimcns Ewunincd: i: soldicrs. \IEXlCO: .falisco, El Rincón, ·1930 lt.
elcv. ·1 soldiers (Coll. ;\ISCollim) VII-17-1968 [Nl\lNH].
NASUTITERMES Dudlcy 1890. Soldiers ol Xa.1utitr:rmes are nasutilorm.
with a conical nasm ami vestigial mandibles with small lmt distinct apict!
points. The shape of thc head behind thc nasus is pyriform, as sccn in
dorsal view. Additionally, for most spccics the head is w.ually clark brown
to black in color. Thc imago head is slightly oval with large protrusive
cyes and occlli. The fontanellc is slit-likc, more-or-les:. anteriorly furcated.
Tibia! spurs are ~:~:~. \Lmdible dentition (Fig. 21'\) ancl wing vcnation
(Fig. 4F) are uscful in gencric recognition. This cosmotropical genu-,
has the most spccies ami the greatest abundance of largc colonies of any
termitc genm. Prohahly much of the 'ucccss of this group is associated
with the cfficacy of the clefcnse strategy of soldiers to squirt glues that art
both mcchanirally ami cbemically against would-be predators, and with
t he adaptation by soldiers to fix atmospheric nitrogen, pcnnitting colonie,
to function cffectively with soldier populations as largc as 30% of the
total nest population. l\csts are frc(¡ucntly arboreal, oftcn in the form of
large brown structurcs made of "carton'', consisting of modified woocl and
sccretions of the workers. A large main nest may be connected with
smallcr suhsidiary nests in nearby trces through a systern of covered nmways, ancl many species do extensivc underground tunnelling. In the
Chamela arca, four species havc becn reportee!, two of which are known
only from ;dates.
Xasutitermcs nigriccps (Haldeman) 1853. The soldiers of Nll.\lltitcrmr.s
11igriu:ps havc dark hrown or hlack hcads with broad conical nasi (Fig.
15E, H) aml the soldier abdominal tergitcs have numerous scatterccl sctac
(Fig. 15B): ;dates are rusty yellow, with clark brown hcads and largc
protrusive compound cycs ;md oce!li (Fig. llG, !-I). Comparcd with ;date"
of pictus (Fig. J(i.\) and colinuu: (Fig. 1GB). 1zigrin:jJs has thc largcst
compound eyc'> (Fig. HiC): in lateral vicw thcy are separated from thc
wntral margin of the hcad by lc>s than (J.~x thc diamctcr o( the
Fig.
~Y.
16. lmago !u
nigrice jJ,\.
eve. The prima:
¡, the most nortl
ging as far nor
northwestern l\Ic
The large colon
to make this sp
posts, ami powe
soldiers cffcctiH
mies. This is thc
Exarnined: 3 in
nest near El Do
?'\ HJ; Chamela.
workers, l win¡
Guenl'ro, Zihu;
l~lR3 [N:\lNH]t
:1 soldiers, R wo
N asutitermes n
distinguishecl in
ml'xicanus na'>t
nigriceps. ~Ior¡
(Fig. l:'íD, H).
The ;date has 1
forested arcas,
on tl-ees. shrut
describcd but i~
:\1<
8)
1is specics are usual! y
comit palms. Obscn·a.nterest, covering such
)lonies; the techniquc.-,
-be predators into daVoucher spccimens of
~reserved. This spccic->
ica, ancl Panama. SfH'EI Rincón, 1 930 IL
NH].
termes are nasutiform,
nall but distinct apicd
s pyriform, as secn in
is mually dark brmrn
with large protrusivc
ess anteriorly furcatccl.
N) ancl wing vcnation
s cosmotropical gcnu-;
f large colonics of any
1is group is associatcd
to squirt glues that act
'e predators, ancl with
en, permitting colonic,
, large as 30% of thc
1, often in the fonn of
of modified wood and
ay be connected with
systern of covercd runld tunnelling. In the
o of which are known
ldicrs of Nasutitcrme~
oacl conical nasi (Fig.
1merous scattercd sct a e
·own heach ancl Llrgc
Compared with alate.,
~riceps has thc largest
re scparatcd f ro m the
the diamctcr of the
KLF
LT (;oLLI'\~.
·¡·ElnllTES OF
IEJ
C!LU!l.L\
B
Fig. !6. llll;]P(}
nigrircjJ.\. "'
.1 . .\"a.llllitermcs
jJiclus;
R.
X.
colimru·;
<..
~V.
eye. The primary quccns become physogastric as the colony ages. This
is the most northcrly-ranging -;pccies of the genm in the Ncw \Vorld, ranging as far north as lower elcvations of Guamúchil (Sinaloa Statc) in
northwcstcrn Mcxico, at lower elcvat ions. lts southcrn cxtension is Brazil.
The large colonics and flexibility in choice of consumccl woocl~ combine
to make this spccies a major pcst of untrcated structural timber, fcnce
posts, ancl powcr lincs. The protective runways and thc numcrous active
soldicr'i effcctivcly detcr ant preclation and attacks by other natural enemies. This is thc only builclcr of large carton nests in the arca. Spccimcns
Ex a m incd: 3 imagoes, 50 solclicrs. 19 "·orkers. 1\IEXICO: jalisco, canon
ncst ncar El Dorado Hotel, 3 soldicrs (Col!. K. 1\[artínC?) VIII-1968 [!\''\[2\'II]; Chamela. on hcach in dcad hollow of living tree. 37 soldicrs, 8
workers. 1 wing. (Col!. D.\l\'ickle ,'\:: \ISCollins ) Vl-29-El8'1 [N\INH]:
Guarcro, Zihuatancjo. 5 soldicrs, 3 workers. (Col!. 1\L Delgado) II-81~lH3 [!\' \1 N 1-I] COI.O\IBIA: ,,úmtcria, in timher roof cciling. 3 imagocs,
5 soldiers, 8 workcrs. (Coll. E. Florcz) VliH-1987 [1\':\INHJ.
1\" asutitermes mexicanus Light l ~133. Soldiers of this species are easily
clistinguished in thc fielcl from solcliers of nigriccjJs on thc basis of color:
11/csicanus na~utcs ILt\·c clark smoky yello"·· much lighter hcads than
nigriceps. \lorphological diffcrences incluclc its long cylinclrical nasus
(Fig. J:,D, H). and sparsity of sctac on the abdominal terga (Fig. IS"\).
Thc ;date has not heen describcd. Colonics o( this spccics are found in
forcstcd areas, building numerous 'lender, light-hrown covcrccl runways
on trccs, shrubs, ami buildings (Light 1~)3?3). Thc nest has uot hccn
dcscribccl but is prohably subtcrrancan ami ccrt;Jinly incompicuous. Some
120
FoLIA E:--;TmroL. ~In.
7'i (1988)
damage to buildings has been ascribed to this terrnite. Light (1933)
suggestccl that the unknown alate may be either colimae or pictus, both
specics clescribecl from alates only from the >ame general rcgion. Rearing
colonies containing nymphs and solclicrs until thc ;date~ cleyclop woulcl
resolve this problem. \Ve urge collectors prcscnt at thc Estación de Chamela betwecn July and September to observe flighb to dt:tcrmine their
sourcc in order to associate the swarmers with soldiers :md young workers.
This is anothcr species with a disjunct di-,tribution. It ¡, found in Colima
and Jalisco S tates on the westcrn coast of ~Iexico, and we collectcd it in
1981 in Veracruz, on thc castcrn coast. Specimrns E.\ m// in NI: 24 soldiers,
35 workers. ~IEXICO: jalisco, Estación de Chamela. UN.\~I. 19 soldiers,
21 workers (Col!. DX"\ickle & l\ISCollins) \'J-2<l-1f)K 1 [~~I~H, ~ICZl:
same data, 5 soldicrs, 1,¡ 1mrkers [:t\1\f:\'H].
Nasutitermes co1imae Light 1933. Thc imago of thi, specie., i, the largest
of the four known species of Xasutitcrmcs from this region. Head width
through thc eyes is nearly 2.0 mm. In side view the compound eye is
scparatecl from the ventral margin of the head capsule by a Iittle lcss
than one-fourth its shorter cliameter (Fig. l6B). Thi'í spccic-. "·as described
by Light (1933) on thc basis of two alates cmerging irom an old orange
trce in Colima, Colima, at 7:00 pm. on .\ugusL :;o 11130. ~o other information i~ availablc.
N asutitennes pictus Light 19:)~1. This is the sma l!est a late of the thrcc:Vasutitcrrncs species. Heacl width is about 1.6:'¡ mm (Fig. l6A). In lateral
Yiew, thc cyes are consiclerably less than onc-fourth the diameter of an cyc
from thc ventral margin of the hcad. The alatcs of thi'i species were collectecl on the same evening at the same tree as thme of colimae. ~o other
information on this species is aYailable.
TENUIROSTRITER:\IES Holmgren 191~. ~a,utiiurm so1dicu, ol this gcnus charactcristically havc hcads that are cleep1y constrictecl bctween the
centcr of the head ancl the bac;c of thc nasus. \Lmdibles are H''itigial, ancl
thc nasth is narrow ancl cdinclrical. The ;mtcnH;tc ha\C l~-1:1 ;¡rtidcs, and
the tibia! ~purs are 2:~:2.
Tcnuirostritcrmes briciae (Snydcr) l ~)~~- This tim aut-likt:, bst-moving
'>pccics 11·;" collected by :\'icklc in a lllüi.-,t, rottcn log- on onc- of thc traih
near Chamcb station. Snyclcr (192~) originally clcscribccl it from material
collectcd in Honduras. The soldicr head. which ¡, Cl'itancous brown, is
much ligl~tcr in color than it is in other spccie> of this gcnm. The head
NICKLE
capsulc is constricted 1
protrusiYc occipital r
1:JC) are clothed witl
bably subterrancan. }
nuirostritcrrncs tcnui
hlack head-;, have ver
of this spccies, one si
mamliblc dcntition (
R, L 1'177. Cat:
Ciencias, Rio de Janein
B\'>KS, K 1906. Two ne''
-----, 1918. Thc tcrmite
'lS: 659-667,
B\'>KS, 1\'. and T. K SN'
notes on biology ancl g·
1-128,
Bt'CKLFY, S, B, 1863. Dese
mol. Soc. Philadelphia ]
Cmu:-:s, M. S. 1969, Wat
of Terrnites, Yo!. l. K.
ami London.
Cou.I:o-;s, M. S. an<l G. D
(Isoptcra, Tcrmitidae):
ciaux 30: 70-81.
Tlrs;-.;Eux, J. 1904. :\o tes
- - - - 1906, YHicties ten
llcm FY, P. H. ami J. B1
Trans . .\'ew York Acad
:\R.\1 JO,
1·Rni:G.HT, W. \\'. 1896.
)J'ales 21: 310-552.
Cmwn, C. ancl J. H.\LL.
imccts from alcohol. Er
II.\Ci :--;, 11. ,\, 1858. 1\[01
Jf.\c¡:,, H. A. 1~60. Mono
fT \LD~l , , . S, S. 1853. TI
Sci. PhiladeljJ!tia 6: 36:
ll\RRI'·· \\'. \'. 1960. Two
1 \('1', 13) j: 253-256.
- ---- _ 19G!. Termitcs, :
It<l. Londnn. 187 pp.
Ho1 ~!CRI::o;, :\. 1911. Ten
Ilaw/1. Bd. 46: 1-86.
EJI 2. Tcrmitcnstuc
.\uensh. l'et. c1had. Han
filll'>'-ü'>, R, A. 1979. Co
of \l'ilrkcr Tcnnitidae
rmite. Light (1933)
imae or pictus, both
teral region. Rearing
tlates dewlop would
.he Estación de Cha' to determine their
i and young 'mrkers.
t is found in Colima
rd we collected it in
;a m ined: 24 solcliers.
Ul\' A~I. 19 soldiers,
81 [:\'~I~H. MCZJ:
species is the largest
region. Head width
1e compound eye is
sule bv a little less
;pecics was described
from an old orange
1930. K o othcr in-
t alate of thc threC'
:Fig. 16A). In lateral
e diameter of an cyc
this species were coof rolimae. :\'o other
n solcliers ol this gcstricted betwcen the
le5 are vestigial. ami
re l~-!3 articles. ancl
mt-likc, fast-moYing
on one of the traih
>ed it from material
:astaneous brown, is
lis genm. The head
l\'rcKLE ET
CoLLI:-~s.
TER:\llTES oF CHA:\IELA
1~1
capsule is constricted behind the antennal sockets, giving rise to an enlarged
protrusive occipital rcgion (Fig. 15F, 1), and the abdominal terga (Fig.
1:"íC) are clothed with numcrous long and short setae. The nest is probab1y subterranean. Although the a1ate is unknown, winged forms of Tenuirostritermcs tcnuirostris, a more northerly species with so1clicrs with
IJ!ack hcacls, have very long, apically curlcd wings. In searching for alates
of this spccies, one should rely on wing size and venation (Fig. "lG) and
mandible dcntition (Fig. 2 (0)).
LITERA TURE CrTED
R. L. 1977. Catalog-o do~ Isoptna do r\ovo Mundo. :\cadcmia Brasilcira ele
Ciencias. Rio de Janeiro. 92 pp.
13\-.:Ks, N. 1906. Two ncw termites. Enlomo[. Nrws 17: 336-337.
~--, 1918. Thc termitcs of Panama ancl British Guiana. Bull A mer ..Uus. Na t. Hist.
'lS: 6'í9-667.
B \'-:KS, N. ami T. E. SNYDER. 1920. A rcvision of the l\'earctic termites (Banks) with
notes on hiology ancl geographic distrihution (Snydcr). U.S . .\'at. Mus Bull. No. 108:
1-128.
Bt:CK!.EY, S. B. 1863. Dcscription of two ncw spccics of tcrmites from Texas. Prnc. Enlomo/. Soc. Philadelphia 1: 212-215.
Coru;o.;s, M. S. 1969. ·water relations in termitcs. Chapter 14. pp. 4~l3-458, in Biologv
of Termites, Yo!. l. K. Krislma and F. M. \Veesner, ccls. Acadcmic Press. New York
and London.
Cm.u-.:s, M ..~. and G. D. PRESTWICH. 1983. Dcfensc in 1\'asutiterrncs octoj>ilis (IIanks)
(Isoptera, Tcrmitidae): comparatÍI'C effectiveness of sohlicr secrctions. lnsectrs Sociaux 30: 70-81.
DrsxEUX, J. 1904. :\'otes tcnnitologiqucs. Ann. Soc. Entomol. Belg. 48: 146-1.51.
----- 1906. \"a ricties tcrmitolog-iques. A nn. Soc. Entamo/. Belg. 49: 336-'l60.
Tlcm rY, P. H. and .J. BEAUMoNT. 1890. Tcrmitcs of the Istbmus of Panama. Part IT.
Trrms. Xew Yorh Acad. Sri. 9: 157-180.
FRncr:.\TT, \V. \\'. 1896 . . \uslralian Tcrmitidac. Part U. Proc. Linn. ,)"oc. New Sollth
Trales 21: 010-552.
CnRDH, C. and J. HALL. 1979. A critica! point clrier use as a mcthod of mounling
insccts from alcohol. Entorno/. News 90: 57-59.
ll\crx, 11. A. 1858. 1\Ionographie der Tcrmitcn. Part 2. Linn. Entorno/. 12: 1-~4::>.
IL\r.n, H. A. 1860. Monographic dcr Terrnitcn (~acbtrag-.) Umz. Entamo!. 14: 100-128.
II\f.D.\u~·. S. S. 18:i3. Termes nig11·rrp.1 from ""'Iexique" clescribed. Pror . . 1cad. Nat.
Sri. Phi/adl'ij;/Jia 6: 36:i.
lf\RRI'·, \\'. \'. 1<J60. T\\'o nc"· tennites of lile family Tcrmitidae . .4nn. Mag. Xat. Hi1t.
1 )cr. 13) :l: 2)3-256.
-·-·--- l~JGI. Tcnnilcs, Their Recognition aJI([ Control. I.ongm~m Cn"Cn antl Co ..
Itrl. London. 187 pp.
Hol.\H.RE:-1, ="· 1!111. Tcrmitenstudicn. 2. Systcmatik der Tcnnitidac. Svcmh. T'cl ..•lhad.
Ilandl. Bd. 4/i: 1-86.
AR.\l'JO,
-- - lfJI~. Tcrmitenstudicn. 3. Svstcmatik dcr Tennitidae. Die Familie 1\Ictatennitida<'.
'''<'!ISk. T'et .. lhad. Hand. Bd. 4G: 1-166.
¡nw;.s.o~, R. A. 1979. Configuration of the cligesti1·c tubc as an aid to identificatiun
of 1\'(>rkcr Tcrmitidac (I,optcra). Sy,l. Enlomo!. 4: 31-38.
FoLIA E:o-;TO\fOL.
:\1 EX. 77 ( 19:-IR)
lütmN.\. K. l'l()J. A gcnnic n·vtston ami phylogenctic study of tlH' famih Kalotcrmitidae (lsoptcra). Bull. Amer. ,1!us. Nat. Hist. 122: 303-10R.
!.HRHLLE, P. A. 1832. Jnsectcs de L\mcriqne Equinoxialc. in Humboldt, R. Obser. Zool.
\Ol. 2. 111.
l.tGHT, S. F. 1927. A new and more exact mcthod of cxpressing importan! spnilic chara<tns of tcnnitcs. Univ. Calif. Pub!. Enlomo[. 4: 75-88.
--- El30a. The California specics of the gcnus A 1nitcnnr'S Sih es tri (lsoptera ). Uuiu.
Calif. Pul1/. Eutomol. 5: 173-21·1.
______ 1930h. Thc Mcxican species of Ami/ames Siln:stri (hoptcra). l'nh•. Calif. Pub/.
F11tomol. r,: 2F,-232.
_____ 19'12. Contrihution toward a rcvision of thc Amnican s¡JCcics of :lmitennn
Silvcs!li (lsoptera). Calif. Univ. Pub/. Enlomo/. 5: :l05--ll1.
_______ I'J:l:l. Teunites of \Ves!em Mexico. l'niv. Calif. Pub/. f-."nto111ol. G: /~J-164.
lt!I.NAFl!S. c. VON. 17:)R. s;·stcma .\'aturc, cdit. 10. \Ol. l. ~2:l PP·
\fi\u.m, F. 1~7~. Bcitrage zur Kcnntniss der Tcrmitcn 1-ll. jenaischc l<'illch. Mt~d ..
Xaturw. 7: 333-3:JR.
:\ICKLF, D. :\. and i\1. S. Cot.u:-.s. 198!l. Kns to thc Kalotermiti<bc of Eastn11 l!nitcd
Sta tes "ith the dcscription of ncw Xr•otennn from Florida (1soptcra). Pror. f:'ntomo/.
Sor. 11'a.1h. 91: 269-2H5.
i\:crnNG, \V. L. 196~J. Distrilmtion ami flights of rarc i\'orth .\mcrican dcscrt tcrmit<''
ol thc gcnns A mitermrs (lsoptcra: Termititlae). Pan-Parifir Flltomo/ogist ·f!i: 320-32'\.
-- 1~~~~~- Tcrmitc flight pcriods: Strategics for prcdator awidance? Sorio!Jiol. 1 (2):
I-11-15C!.
Roo:\WAI.L. ~-1. L a!l(l O. H. CtJOTA:\1. 1962. Jndian spccics of the tcrmite gcnns Coj,fotniiU'S En tomo l. i\lonog. :\o. 2. Jndian C:onncil A gr. Res. :\c\1· Dclhi: 11 'i pp .
.~<:I!FFFIL\11:--J, R. H .. L. K. CAsTo~, J. J. SIMS an<l M. K. Rtsr. 1983. l<lentification of
thc ddcmive sccrction from soldiers of the 1\'orth American tcrmitc A miterme.,
u•hn-/cri (llcsncux) (1 soptcra: Tcrmitidae). ] . Che m. Frol. 9: 12~l3-1305 .
.~CIJEFR\11:-\, R. JI.. J. J. StMs. R. K. LEE, a!l(l M. K. Rtsr. ¡<¡¡;(í_ llclminthogermacrcne. a majen romponent in thc dcfensive sccrction of tlw :\carctic termitc. A mitermes wheeleri. 1 .Nat. Products 49: 699-701.
Sli.\'LSTIU, F. EIUI. :\ota prcliminarc sui Tcnuitidi sud-amcricani. Bol/. ,Uns. loo! ..4nat.
ComjJ. Tori1zo IG: 1-8.
~l'\YDFR, T. E. 1922. :\cw tcrmitcs from H;nvaii, Central and Sonth .-\!IHTÍ<a. ancl thc
Antillcs. J'ror·. U.S. ;'\'(¡[. ,1/u.\. 61: l-:l2.
----- 1923. Thrce new tcrmitcs from the Canal Zo11c. Panama, /'roe. Enlo111ol. Sor_
Wa.1h. 2:í: 126-1cll.
------ 1921. Dcscriptions of ncw spccics and hithcrto unkno\\'11 <a.stcs of tcrmitcs frmn
Amcrica and 1-Lm·aii. Proc. F.S. ;\'at. Mus. 64: 1-40.
!'HG. A small dark-colorcd nc\1· Kalot<-r,nr·, from Guatemala. Pror. Entonwl.
Sor. J!'a.1h. 4f': 15~-160.
1'l 19. Ca tcdog o[ !he termitcs (lsoptcra) of thc world. S111iiÍISOIÚ<lll .1Ii1c. Collcrt.
11~: 1 <119.
\\'ALKLR, F. IS'í3. List of !he spccimens of tH'uropterous insccls in thc collcclinn of tlw
British .\Iuscum. l'art. 3, Termitides: ;)01-529.
\\ ASMAl'>N, E. IS'Hi. :\cuc Tcrmitophilcn llttd Termitcn atts Imlicn. \'iaggo di Leonardo
Fea in Hinnattia e Rcgione \'icinc. Ann. del Jf111. Cil·. rli Stor. Xat. (;eno;•a IG:
Gl3-630.
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